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Issued by
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. U.S.A.
APRIL 1996
ATOLL RESEARCH BULLETIN NOS. 435-442
RESEARCH
BULLETIN
ATOLL RESEARCH BULLETIN
NOS. 435-442
NO. 435.
NO.
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442.
MORPHOLOGY AND MARINE HABITATS OF TWO
SOUTHWESTERN CARIBBEAN ATOLLS: ALBUQUERQUE AND
COURTOWN
BY JUAN M. DIAZ, JUAN A. SANCHEZ, SVEN ZEA, AND
JAIME GARZON-FERREIRA
CORAL FAUNA OF TAIPING ISLAND (ITU ABA ISLAND) IN
THE SPRATLYS OF THE SOUTH CHINA SEA
BY CHANG-FENG DAI AND TUNG-YUNG FAN
FIRST OBSERVATIONS ON THE FISH COMMUNITIES OF
FRINGING REEFS IN THE REGION OF MAUMERE (FLORES-
INDONESIA)
BY MICHEL KULBICKI
GROUPER DENSITY AND DIVERSITY AT TWO SITES IN THE
REPUBLIC OF MALDIVES
BY ROBERT D. SLUKA AND NORMAN REICHENBACH
EFFECT OF TYPHOONS ON THE LIZARD COMMUNITY OF A
SHELF ATOLL
BY MICHAEL JAMES MCCOID
FLOWERING AND FRUITING IN THE FLORA OF HERON
ISLAND, GREAT BARRIER REEF, AUSTRALIA
BY R.W. ROGERS
NAMU ATOLL REVISITED: A FOLLOW-UP STUDY OF 25
YEARS OF RESOURCE USE
BY NANCY J. POLLOCK
CRUSTACEA DECAPODA OF FRENCH POLYNESIA
(ASTACIDEA, PALINURIDEA, ANOMURA, BRACHYURA)
BY JOSEPH POUPIN
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
APRIL 1996
ACKNOWLEDGMENT
The Atoll Research Bulletin is issued by the Smithsonian Institution to provide an
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that supports the biota. The Bulletin is supported by the National Museum of Natural
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COORDINATING EDITOR
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ATOLL RESEARCH BULLETIN
NO. 435
MORPHOLOGY AND MARINE HABITATS OF TWO SOUTHWESTERN
CARIBBEAN ATOLLS: ALBUQUERQUE AND COURTOWN
BY
JUAN M. DIAZ, JUAN A. SANCHEZ, SVEN ZEA, AND
JAIME GARZON-FERREIRA
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ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
APRIL 1996
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MORPHOLOGY AND MARINE HABITATS OF TWO SOUTHWESTERN
CARIBBEAN ATOLLS: ALBUQUERQUE AND COURTOWN
BY
JUAN M. DIAZ}, JUAN A. SANCHEZ , SVEN ZEA!” and
JAIME GARZON-FERREIRA'
ABSTRACT
Albuquerque and Courtown are two small, uninhabited oceanic atolls, located in
the southwestern Caribbean Sea, belonging to the San Andrés and Providencia
archipelago, Colombia. These atolls have a volcanic basement and are surrounded by deep
water. Based on photo-interpretation of geomorphological and ecological features as well
as on data collected during field work, the gross morphology, marine bottom habitats and
reef structures of both atolls are described down to a depth of 50 m. Distributions of
morphological and bottom habitat units are presented in thematic maps showing the
overall zonational patterns in the two atolls.
Morphological and ecological zonations in both atolls are primarily controlled by
both wave exposure in a windward-leeward gradient and depth. The presence of an ample
windward fore-reef terrace, a well developed windward barrier reef with spur-and-groove
system, an extensive lagoonal terrace with sudden transition to the lagoon basin, and
profuse development of ribbon and anastomosing patch reefs in the lagoon are
characteristics common to both atolls. As in other Caribbean atolls, the outer slope in
Albuquerque and Courtown is outlined by a sandy step or bench at 35 to 45 m depth.
Significant differences between the two atolls exist in the degree of development and
structure of leeward peripheral reefs, as well as in the amplitude of the leeward fore-reef
terrace. At Albuquerque, peripheral reefs grow on a shallow flat and enclose the lagoon
along a wide semicircle, whereas at Courtown such reefs have in part developed algal
ridge-like structures and are unevenly distributed, leading to an open lagoon to the east.
The broad leeward terrace in Albuquerque contrasts markedly with the rapidly dipping
leeward slope towards the outer shelf margin in Courtown.
Accumulations of sand and rubble have led to the formation of cays and small
islands on the lagoonal terrace in both atolls, but also on leeward peripheral reefs in
Instituto de Investigaciones Marinas y Costeras, INVEMAR, Apartado 1016, Santa Marta,
Colombia
* Universidad Nacional de Colombia (Departamento de Biologia)
Manuscript received 31 March 1995; revised 21 Novermber 1995
Courtown, some of which have experienced remarkable changes in their size and shape in
the last 25 years.
Biological composition and structure of reefs in both atolls show a great
resemblance to one another and to the better-known reef complexes around the nearby
islands of San Andrés and Providencia. Although no urban development exists in these
atolls, recent decline of living coral and over-exploitation of marine resources were
evident.
INTRODUCTION
There are about 425 atolls worldwide and only 15 of them are located in the
Atlantic, of which four are part of the San Andrés and Providencia archipelago in the
Southwestern Caribbean Sea (Milliman, 1973; Geister, 1983). This archipelago comprises
a series of islands, atolls and coral shoals running in SSE-NNE direction, parallel to the
Nicaraguan Rise for more than 500 km. It is separated from the Central American
continental shelf by the San Andrés Trough (Fig. 1). The southernmost reefs of the
archipelago, Albuquerque and Courtown (the latter are also called Bolivar Cays) are two
small atolls lying about 200 km east of the Nicaraguan coast. Although geographically
closer to Central America than to the South American continent, the archipelago has
belonged to the Republic of Colombia since 1822.
The accurate date of human discovery of these atolls is uncertain but their
locations were well known to the Spanish sailors of the 16th century and were probably
occasionally visited by Miskito Indians from the Central American coast, who came for
fishing and turtling (Parsons, 1956). None of the tiny sand cays on the atolls has sufficient
land to warrant permanent settlement, but one of them on each atoll serves presently as a
military post for the Colombian navy, and they are visited regularly by fishermen and
tourists in chartered yachts from nearby San Andrés.
Briefly mentioned by Darwin (1842) in his interpretation of Caribbean reef
structures and their origin, the reefs of Albuquerque and Courtown have since received
little scientific attention in comparison with those around the nearby islands of San Andrés
and Old Providence (Geister, 1969, 1973,1975,1992; Kocurko, 1977; Marquez, 1987;
Diaz et al., 1995) and other West Atlantic and Caribbean atolls such as Hogsty Reef
(Milliman, 1967), Alacran Reef (Kornicker and Boyd, 1962; Bonet, 1967), Chinchorro
(Jordan and Martin, 1987) and those off Belize (Stoddart, 1962; James and Ginsburg,
1979: Riitzler and Mcintyre, 1982; Gischler, 1994). Albuquerque and Courtown were
briefly visited by the Fifth George Vanderbilt Expedition in 1941. Published observations
include reports on the birds (Bond and DeSchauensee, 1944), fishes (Fowler, 1944) and
crustaceans (Coventry, 1944). The R/V GERDA, of the University of Miami, stopped in
May 1966 for few days at Albuquerque and Courtown and conducted observations on the
ecology, morphology and oceanography of the atolls. From this visit, Milliman and Supko
(1968) made preliminary conclusions on the geological origin, and Milliman (1969)
described the general characteristics of the reefs and commented on hydrography. Further
oceanographic findings from the waters surrounding the atolls have been recorded during
research cruises by the Colombian navy (Gonzalez, 1988; Téllez et al., 1988). Aspects of
the terrestrial environment and fauna were more recently discussed by Chirivi (1988).
However, very little is known about the distribution of marine bottom habitats and the
zonation of the reefs constituting these atolls.
Therefore, the purpose of this paper is to give the first detailed systematic
description of the gross morphology and the marine habitats of Albuquerque and
Courtown atolls, with emphasis on the reef structures.
REGIONAL SETTING
Albuquerque and Courtown are the southernmost reef complexes of the San
Andrés and Providencia archipelago. Albuquerque (12° 10' N and 81° 51' W) is located
37 km south of San Andrés Island and about 190 km east of the Nicaraguan coast. It is
nearly circular in shape, about 5.5 km E-W and 4.5 km N-S. Two small islands, North
Cay and South Cay, rise up to 2 m above mean sea level behind the seaward barrier reef
and are separated from each other by a 250 m shallow channel.
Courtown (12° 24' N and 81° 28' W) lies 30 km southeastward from San Andrés
and 47 km northeast of Albuquerque. It is kidney shaped, about 3.5 km E-W and 6.5 km
SSE-NNW (Fig. 3). Although this atoll presently bears two cays (East Cay and West or
Bolivar Cay) and a tiny sand spit, their size, shape and number seem to be quite variable in
the course of time, as can be easily inferred from an earlier description and map of the
atoll by Milliman (1969).
Toward the north and eastern sides of both atolls, an almost continuous barrier
reef is well developed, whereas the leeward peripheral reefs are absent or ill defined and
are separated by wide gaps and channels.
Both atoll foundations rise from the surrounding sea floor more than 1000 m deep,
and apparently have a volcanic basement. Unequivocal evidence for the volcanic origin of
these atolls and nearby islands comes from the magnetic anomalies detected at San Andrés
Island and Courtown, one volcanic pebble dredged from Albuquerque basement (Milliman
and Supko, 1969), as well as the volcanic rocks of Providencia (Geister, 1992) and the
Corn Islands (McBirney and Williams, 1965). Further aspects of the geological origin of
the archipelago are discussed by Geister (1992: p. 56-58)
Available meteorological data recorded from nearby San Andrés between 1959
and 1986 (Diaz et al., 1995) are used here, as there are no recorded observations from
either atoll. The mean annual air temperature is 27.4°C, with a 1°C range in monthly
values. The annual rainfall measured at San Andrés is about 1900 mm, of which over 80%
falls between June and November. Winds are trades, from the ENE, with a mean annual
intensity of 6.1 m/s and mean monthly variations between 4.5 m/s (May, September-
October) and 6.6 m/s (December-January, July). Sporadic storms occur mostly in the
second half of the year, with westerlies or northwesterlies attaining speeds over 20 m/s.
Albuquerque and Courtown lie in the Caribbean hurricane belt. Hurricanes were
recorded in 1818, 1876, 1877, 1906, 1940, 1961, 1971 and 1988 (cf. Barriga et al., 1969:
23; Geister, 1992: 7; Diaz et al., 1995: 112). The latter, 'Joan', on October 20-22 1988,
passed westwards 90 km south of San Andrés (about 50 km south of Albuquerque); its
eye attained a diameter of about 35 km and the wind reached speeds over 210 km/h
(Geister, 1992).
The Caribbean Current reaches Albuquerque and Courtown from the NE with
speeds of 0.5-1 m/s and passes over the atoll shelf in a SW to W direction, being highly
affected by the irregular bottom topography of the shallow-water zones. Waves are
generated by the trade winds and approach the atolls from the NE to E, the effective fetch
extending for nearly 2,000 km over almost the entire width of the Caribbean Sea. Hence,
the considerable amplitude and height of waves breaking on the barrier reef along the
windward side of the atolls.
The sea surface temperature averages 27.5°C, with mean monthly values ranging
between 26.8 (February-March) and 30.2°C (September-October). Surface salinity
fluctuates between 34.0 and 36.39/00 (Gonzalez, 1988). Tides on the atolls are mixed
with a strong diurnal component. Tidal ranges between 0.3 and 0.6 m are recorded from
nearby San Andrés (Geister, 1975).
METHODS
A preliminary photo interpretation of geomorphological and ecological features of
both atolls was done on panchromatic total coverage air photography taken in 1971 and
1984 by the Colombian Geographical Institute (Instituto Geografico 'Agustin Codazzi')
approximately 1:22,500, 1:23,000 and 1:30,000, which was then used as basis for field
sampling. Preliminary morphological and habitat distribution maps at 1:20,000 scale were
drawn combining reef and lagoon photo-patterns defined on the basis of tone, texture and
location, as well as bottom topography inferred from bathymetric charts 1:20,000 COL-
203 (Albuquerque) and COL-204 (Courtown). Further detail of the spur-and-grove
system of the barrier reefs and lagoonal patch reefs was obtained from oblique aerial
colour slides taken on September 29, 1994 from a chartered aircraft at altitudes of 200 to
500 m.
During a cruise to the atolls in May-June 1994 aboard the R/V ANCON of the
Instituto de Investigaciones Marinas y Costeras (Santa Marta, Colombia), 8 days (May
20-27) were spent at Courtown and 12 (May 28 - June 8) at Albuquerque. A total of 23
(Courtown) and 25 (Albuquerque) observation and sample sites were visited (Figs. 2-3).
Location of sample sites included several examples of each of the photo-pattern units, and
their exact geographical placement was carried out with an accuracy of 20 m with the aid
of a portable Geographic Positioning System (GPS) instrument. SCUBA was used for
depths over about 6 m SCUBA was used, otherwise observations were made while skin
diving or walking for shallower areas. Observations of bottom types, depth, direction of
currents, dimensions and distribution patterns of the reef structures, as well as species
composition of dominant biota were recorded on acrylic data sheets. Complementary
depth profiles were recorded with the ship's echosounder (28 khz).
Final thematic maps at 1:20,000 (bathymetry, geomorphology, bottom habitats,
wave exposure) were entered via a digitizing table into a geographic information system
(GIS-ILWIS) for storage, processing and further analysis. Morphology and _ habitat
classification and terminology vary considerably between authors, and the terms used here
to define morphological units and reef zones follow those of Geister (1975, 1977, 1983).
Marine habitats are named, where possible, after the substrate dominating macrobiota or
substrate features, as was done by Duyl (1985) for the reef environments of the
Netherland Antilles.
RESULTS
Both atolls have the same basic morphological features (Figs. 2 and 3) and, with
minor differences, the same marine environments (Figs. 4 to 7). To save space, a general
description of each of the morphological units is given below with comments on the
bottom habitats found there (map units on Figs. 6 and 7) and, where necessary, on the
pecularities of each atoll. Table 1 includes a brief description of the habitats (map units)
and allows cross referencing to morphological units. Figures 4 and 5 are representative
profiles of the atolls and show the morphological features and bottom habitat distribution
along a windward-leeward (right to left) gradient.
FORE-REEF TERRACE AND OUTER SLOPE
The windward margin of the atolls is characterized by the presence of a gently
dipping terrace, descending at low angle (from 6 to 9 degrees) to -24 to -30 m (somewhat
deeper in Albuquerque than in Courtown), where a topographical break gives way to a
subvertical slope below -30 m. The break marks the transition to the outer slope of the
atoll shelf. From a depth of 4-8 m seaward of the barrier reef, to about -15 m, this flat,
calcareous platform is, with the exception of scattered gorgonians (Pseudopterogorgia
sp.) and large sheets of excavating sponges (Cliona aprica and C. caribbea), mostly
devoid of sessile organisms and sediments (‘bare calcareous hard bottom' unit). Low relief
calcareous ridge-like structures, with a parallel layout similar to the spur and groove
system of the barrier reef (see below), are found along the entire width of the terrace and
are more conspicuous on the northeastern section of the atolls. Shallow furrows between
these low ridges are filled with coarse sediments and rubble below 18 m. Toward the
outer margin of the fore reef terrace, faunal richness and diversity increase gradually, at
first especially with brown algae (Sargassum sp., Stypopodium sp.), green algae
(Halimeda spp.), massive scleractinians (Diploria spp., Porites astreoides, Siderastrea
siderea) and many branching octocorals (Pseudopterogorgia spp., Pterogorgia citrina,
Eunicea spp., Plexaurella spp.) (‘Gorgonaceans on hard bottom’ unit). Below 18 m more
and more hemispherical scleractinians (Montastraea spp., Colpophyllia natans, and
others) and sponges come into sight, as well as coarse sediments that accumulate in
shallow hollows. Although coral heads often attain considerable size, they are mostly
solitarily, tens of meters apart. In contrast to this, a narrow belt along the transition zone
to the outer slope (24 to 30 m) exhibits a well developed coral community, and the
calcareous platform appears therefore almost totally covered by corals (Vontastraea spp.,
Colpophyllia natans, Agaricia agaricites, Dichocoenia stokesii, Stephanocoenia
intersepta, among others) , algae (Lobophora sp., Halimeda spp.), sponges and
octocorals (‘mixed corals’ unit).
The windward outer slope was visited only in Courtown, but its morphology
seems to be similar in both atolls, as could be inferred from the recorded bathymetric
profiles. The outer slope dips gently (ca. 40-50°) to a sand step beginning at -30 to -35 m.
Since this step can be easily recognized on the aerial photographs as a narrow, light grey
band along the windward margin of the atolls, thus it seems to be covered by high-
reflectance sediments. Below this step, the outer slope decreases subvertically to -400 m
and then at lower angle to depths beyond 1,000 m.
WINDWARD BARRIER REEF
The barrier reef does not completely encircle the atolls, but extends only along the
inner shelf from the NNW, N, NE, E, and SE almost continuously for about 5.6 km at
Albuquerque and 7.5 km at Courtown. The continuous reef segments are 50-250 m
across, being formed by more or less coherent ridges rising from the upper margin of the
fore-reef terrace at 5 to 6 m to a reef flat near low tide level.
The barrier reef is normally deeply penetrated by surge channels oriented
perpendicular to the reef front, forming a typical spur-and-groove system which is easily
recognizable on the aerial photographs. Also scattered coral pinnacles rise in some places
from about -4 m, just windward of the surf zone, often breaking the surface. The spurs
rise 0.5 to 2 m above the adjacent grooves, the latter being 1 to 5 m or more wide and
often exhibiting anastomosing bifurcations (Plate 1). At Courtown, the barrier reef is
indented at two places, giving the atoll its distinctive kidney shape. Here, the reef crest
becomes discontinuous, and a well developed buttress-groove system appears instead
(Plate 2), creating a transition zone 300 to 500 m wide between the fore-reef terrace and
the lagoonal terrace in its lee. The 2-3 m depth surge channels in this area allow small
boats to pass the barrier during calm days. At Albuquerque, the barrier reef is virtually
continuous, but on its NE margin a few unusually wide grooves interrupt the reef flat for
10 to 20 m, permitting some waves to pass undisturbed into the lagoonal terrace. At this
place, a second, discontinuous barrier reef, located 100 to 200 m behind the former and
nearly parallel to it, generates a displaced surf zone clearly observable from the air.
The main framework builder in the windward reef flat is the hydrocoral Millepora
complanata, which is commonly associated with incrustations of coralline algae.
Millepora and the zoanthid Palythoa sp. overgrow the shallowest zone of the barrier reef
flat and the upper surfaces of the spurs (‘Millepora-Palythoa' unit), the high surf splashing
and washing permanently the emergent colonies. In the buttress-groove area in Courtown,
as well as in the second barrier at the NE margin of Albuquerque, Palythoa is generally
replaced by Porites porites (growing usually within the Millepora colonies) and crustose
forms of Porites astreoides and Diploria clivosa, which overgrow with Millepora the
upper parts of the buttresses and the reef flat (Willepora-P. porites' unit). The upright
sides of the spurs and buttresses are encrusted with Diploria spp., Porites astreoides and
Agaricia agaricites, often assuming a flat form. The hydrocoral Stylaster roseus, the
green alga Halimeda, as well as coralline red algae (Porolithon sp.) are also common
elements in this zone. Large (up to 2-3 m in diameter) sheet-like excavating sponges
(Cliona aprica, C. caribbea) may be fairly common at the sides and bottom of grooves.
Leeward of the reef crest, following the ‘Millepora-Palythoa’ unit, cushion-like
colonies of Porites porites as well as massive P. astreoides and Diploria strigosa occur at
some places among small ridges of Millepora and calcareous boulders (‘Millepora-P.
porites’ unit). The displaced rear barrier reef on the NE side of Albuquerque consists
likewise of extensive ridges with Millepora complanata and Porites porites rising from -
1.5 to -2.5 m. In some places, like in the NE barrier of Albuquerque and the SE section of
Courtown, the coral growth on the rear reef zone extends for about 250 m. There, the end
of the barrier reef is marked lagoowards by patchy thickets of Acropora palmata,
accompanied by small colonies of Diploria strigosa, Montastraea spp. and occasionally
also by cushion shaped colonies of Porites porites (‘Diploria-A. palmata’ unit).
In Courtown, the southernmost portion of the barrier reef becomes discontinuous
after it bends westward. Numerous pinnacles, constituted mostly by a framework of
Millepora at their upper parts, rise in this area from -4 to -5 m reaching up usually to a
few centimeters below the surface (Plate 3). At the base of the pinnacles are massive
colonies of Diploria spp. commonly more than 2 m in diameter, small thickets of
Acropora cervicornis and branching octocorals. The pinnacles are generally arranged in
groups, separated by anastomosing sandy channels, with a characteristic wave-induced
pattern of ripple marks. Coral rubble (mostly of Acropora cervicornis) accumulates at the
sides of the channels.
LAGOONAL TERRACE
The leeward margin of the reef flat leads down to the lagoonal terrace usually with
an abrupt, 0.6 to 1.5 m high, steep slope. The lagoonal terrace is a flat platform attaining a
width of 200 to 900 m and increasing in depth from 1 to 3 m towards its inner margin.
The lagoonal terrace is one of the most discernible features from the air due to its pale
hue. Close to the rear reef, the terrace is covered by rubble (‘hard bottom and rubble'
unit), which is gradually replaced lagoonwards by gravel and coarse sand (‘sand and
rubble’ unit). The rubble zones are usually arranged in elongated layers perpendicular to
the barrier reef, apparently related to the grooves and depressions of the reef crest. The
innermost rubble areas on the terrace are overgrown by green (Halimeda, Padina), brown
(Dictyota, Turbinaria) and red algae (Amphiroa, Neogoniolithon), as well as scattered
encrusting scleractinians (P. astreoides, Siderastrea)('‘rubble with algae’ unit). Some
portions of the sandy bottom, particularly in Albuquerque, are sparsely colonized by green
algae (Penicillus, Rhipocephalus, Udotea), where juvenile individuals of the gastropod
Strombus gigas are fairly common. The lagoonal terrace normally terminates on its lee
with a steep 'sand cliff, leading down into the lagoon basin with slopes up to 40°. It
represents an accretionary fore-set of fine-grained sediments transported from the reef
area to the leeward margin of the terrace
The two cays existing in Albuquerque (North Cay and South Cay, Plate 4), as well
as East Cay in Courtown (Plates 5 and 6), are sand and rubble accumulations on the
lagoonal terrace. Coconut palms, Ficus trees, Scaevola bushes and Tournefortia shrubs
are the dominant vegetation. North Cay, at Albuquerque, serves today as military post for
the Colombian navy. Several bands of beachrock, paralleling the windward shoreline of
these cays, extend eastward on the lagoonal terrace for about 15 (both cays in
Albuquerque, Plate 7) to 70 m (East Cay in Courtown), suggesting the location of
previous shorelines and thus a lagoonward migration of the cays. The two cays at
Albuquerque are presently very close to the leeward margin of the lagoonal terrace.
Where submerged beachrock is not covered by rubble and sand, it is mostly overgrown by
encrusting coralline and green algae (Halimeda, Rhipocephalus) that contain dense
populations of boring sea urchins (Echinometra lucunter). The only sea grasses on the
atolls occur on the sheltered leeward side of North Cay in Albuquerque and East Cay in
Courtown, where they cover the shallow sandy bottom of the terrace (‘sea grass' unit) The
dominant grasses in Courtown are Syringodium and Halodule, whereas Thalassia is more
abundant in Albuquerque. The edible urchin, 7ripneustes ventricosus, is abundant in these
grass meadows.
LAGOON WITH PATCH REEFS
The depth of the lagoonal basin is as much as 18 m (in Albuquerque, see below)
but generally it varies between 8 and 10 m. Where corals and coral reefs are lacking, the
lagoon floor is covered by white calcareous sediments, the coarser fractions of which
consist mostly of fragments of coral, molluscs, foraminifera, coralline algae and Halimeda,
and rubble. Numerous burrows, mouds and faecal pellets throughout the deeper parts of
the lagoon evidence an active bioturbation of the bottom (‘bioturbated sediments’ unit).
Green algae (Rhipocephalus, Udotea, Halimeda) grow sparsely around coralline areas
forming small patches, where one or more individuals of the Queen Conch, Strombus
gigas, as well as patchy aggregations of garden eels (7aeniconger sp.) are occasionally
found.
A significant portion of the lagoon is occupied by coral reefs, which are highly
variable in shape and size, as well as in the dominant scleractinian species, depending
mainly on the depth and wave exposure. Reefs occur as solitary mounds and miniatolls, or
as ribbon and anastomosing patch reefs. In order to simplify the nomenclature, we divided
the patch reefs found inside the lagoon into three main types (map units, see Table 1),
according to the dominant scleractinian species: a) emergent to very shallow 'Diploria-A.
palmata’ reefs dominated at their summit by Diploria strigosa and Acropora palmata, b)
2-5 m deep 'A.cervicornis' reefs dominated by thickets of Acropora cervicornis, and c) 4-
16 m deep '‘Montastraea spp'.-reefs dominated by one or more species of the Montastraea
annularis species complex (see Weil and Knowlton, 1994).
At Courtown, lagoon depths vary between 7 and 15 m. Patch reefs cover about
30% of the lagoon floor. In the northern half of the lagoon, where the average depth is
about 10 m, a dense net of anastomosing reefs (Montastraea spp.) covers nearly 50% of
the bottom. Most of them are low-lying, rising no more than 4 m above the bottom (Plate
8), but some are nearly emergent and form a wave-breaking zone of thickets of A.
palmata. The relative coverage of living scleractinians composing these reefs ranges
between 10 to 50% from one patch to another. In many places, heads of Montastraea
annularis are extent overgrown by filamentous and brown algae (mainly Lobophora
variegata), and scattered thickets of Acropora cervicornis are up to 90% devoid of living
tissue. Although the bottom in the central and southern portions of the lagoon is
predominantly covered by sand, solitary mounds and scattered coral heads are common.
In some places of the central area, large aggregations of single coral heads and small
thickets of A. cervicornis occur (at present largely dead), forming diffuse, non-cohesive
reef communities. The lagoon is rather open to the E and NE, lacking a well defined sill.
Nearly 25% of the lagoon floor at Albuquerque is covered by patch reefs. The
lagoon exhibits two distinctly depth levels, which are easily recognized from the air
because of their different blue hues (Plate 9). A first level, with an average almost
constant depth of 9 m, takes up the N and E parts of the lagoon and about 65% of its
whole area. The second depth level averages about 15 m and takes up the leeward half of
the lagoon to the W and S. Both levels are separated by a meandering ribbon reef of
'‘Montastraea spp.', which wanders for nearly 6 km, attains 10 to 30 m in width and rises
up to -4 m. On the upper lagoon level there are also several nearly circular shaped
miniature atolls which break the surface. These reefs are of type Montastraea spp. at their
base but show a typical zonation to the 'Diploria-A.palmata' type towards the summit.
Anastomosing patch reefs (Montastraea spp.), with the same basic structure as those at
Courtown, are found in the northeastern and southeastern parts of this lagoon level. The
deeper level is more sparsely covered by reefs. These are mostly low-lying, isolated patch
reefs of the ‘Vontastraea spp' type. The depth of the lagoon diminishes leewards to about
-5 m or less and the bioturbated sediments of the bottom give way to a gravel-rubble
zone, representing the lagoon sill and the transition zone to the western terrace.
According to our observations, lagoonal currents are completely wind-driven and
perceptible over the entire water column. Although some differences in direction and
10
intensity were noticed from one location to another, average current velocities of about
2.5 m/min were estimated on the surface at an almost constant wind intensity of 3 m/s in
Courtown, and of about 3.5 m/min (wind velocity: 6.5 m/s) in Albuquerque. Considering
the rather small size as well as the shallow and open nature of the lagoons, the residence
time of lagoonal water masses are thus apparently short, probably not exceeding 24-36
hours.
LEEWARD PERIPHERAL REEFS
Leeward peripheral reefs are poorly developed in both atolls. In Courtown, the
absence of such reefs for more than 2 km results in a widely open lagoon to the west. The
northernmost portion of the barrier reef becomes interrupted after it curves
southwestward semi-enclosing the northern part of the lagoonal terrace. Southwards,
detached reef flats rise from 5 to 7 m depth and break the surface in irregular intervals of
50 to 400 m for about 1.4 km, building the northern peripheral reefs. Wave refraction
around the north end of the atoll results in colliding surf from both the NE and the NW.
Similarly, beginning at the southwestern tip of the atoll, a series of detached reefs and
Shoals semi-enclose the southern third of the lagoon. Some of these reefs are partly
emergent at low tide and most of them are almost completely coated by calcareous red
algae (Porolithon sp., 'coralline algae! unit), resembling the algal ridges characteristic of
Pacific atolls. The algal crust usually exhibits numerous bores caused by chitons
(Choneplax lata) similar to the systems described elsewhere in the Caribbean (Littler ez
al., 1995). Scattered colonies of Diploria strigosa and Millepora encrust the reef flat,
whereas on the subvertical to overhanging walls Dendrogyra cylindrus, Agaricia
agaricites, branching octocorals (Plexaura sp., Pseudoplexaura sp.) and bunches of
Halimeda are common. Wave turbulence, swift currents and the presence of an intricate
system of caves in the northern and southernmost peripheral reefs in Courtown create a
bizarre and attractive environment. Sand and rubble accumulations over the larger leeward
peripheral shoals at Courtown led to the formation of one island (formerly two, see
discussion) and a small sand spit. The island serves today as military post for the
Colombian navy (Cayo Bolivar).
At Albuquerque, leeward peripheral reefs grow on a shallow, wide sand flat,
which represents the lagoon sill. A series of small, low lying reefs enclose the lagoon basin
along a wide semicircle between the northwestern tip of the barrier reef and the southern
margin of the lagoon. Two navigable channels on the NW and SW breach the flat into the
lagoon basin. The peripheral reefs are constituted mainly by large thickets of Acropora
palmata, as well as isolated heads of Diploria strigosa and Porites astreoides. Crustose
coralline algae (Porolithon sp.), coating large areas of the coral framework, are also major
constituents of these reefs. Octocorals and dense beds of brown algae (Dictyota)
extensively cover the reef flat bottom. In some places, the scleractinians are dead and
overgrown by Dictyota or encrusted by coralline algae. The patch reefs in the
southwestern edge, at both sides of the navigable channel, are particularly affected. Here,
large thickets of A. palmata were found broken and even overthrown. Large amounts of
coral debris were dispersed around the reef flat, including fragments of A. cervicornis, at
present an uncommon species in Albuquerque's reefs. This perturbation may have been
11
caused by hurricane ‘Joan’, whose eye passed westward in October 1988 only a few
kilometers south of Albuquerque, with winds of more than 200 km/h, which produced
very abrasive swells from the south.
LEEWARD TERRACE AND OUTER SLOPE
In the leeward margin of Courtown Atoll there is not a well defined fore-reef
terrace. A slope descends in a distance of no more than 200 to 300 m from the shallow
reef flat or the lagoon sill to 17 to 20 m, giving rise suddenly to a subvertical sand slope or
to a vertical cliff with locally overhanging ledges. The sand-covered slope of the terrace
acts as sedimentary ramp, across which reef detritus falls to greater depths. In the
northern and central sections, extensive but somewhat diffuse coral carpets cover as much
of the bottom, forming elongated low buttresses in an E-W direction and alternating with
rather broad sandy channels. Much of the coral (ca. 75% of the bottom) is at present dead
and overgrown by fleshy brown algae (Lobophora, Dictyota), whereas living
scleractinians cover no more than 10% of the bottom. Although in the southern half of the
terrace coral carpets are scantier and have a patchy distribution, they are better developed
and form a distinct hardground on the sandy slope, showing a coverage of nearly 70% of
living tissue (‘scattered corals’ unit).
At the outer edge of the terrace, the angle of the sandy slope increases to nearly
45°, whereas the reef slope drastically changes to a near vertical wall at about -15 m.
Species richness and abundance of scleractinians are very high on the outer margin of the
terrace, where massive Montastraea annularis, M. franksi, M. cavernosa and
Colpophyllia natans form especially in the southern part, large dome-like structures rising
up to 3 m above the bottom (‘mixed corals’ unit). Between these structures usually run
‘sand rivers', which continue as sand falls on overhanging locations along the drop-off.
Apart from scattered, small plate-like agariciids such as Agaricia undata, the vertical cliff
is mostly devoid of corals and the only organisms attached to the rather smooth
substratum are large tube-like and ramose sponges (Agelas conifera, Aplysina spp.,
Totrochota birotulata), antipatharians and clumps of Halimeda. At the southern locality
visited, the cliff remains vertical to about -45 m, where a slanting sand-covered step,
about 40 m wide, lines the outer slope of the atoll shelf. In this area, the sand-covered
bench deepens at an angle of nearly 30° to about -55 m, where a steep slope continues to
greater depths. The loose sand on the slope is composed of Halimeda with accessory shell
and coral grains. Large plate-like corals (probably Agaricia and Montastraea) and
antipatharians could be observed from above growing along the outer margin of the sandy
slope. At another locality, situated in the central section, the drop-off is subvertical to -28
m and is mostly covered by plate-like scleractinians (Agaricia, Montastraea), the sand
step is much wider and dips at a lower angle. It seems probable that such a sandy step
does occur along the entire leeward margin of the atoll, although the indicative lighter
photo-pattern is not always visible on the aerial photographs, possibly due to its variable
slope angle and width.
In contrasting to Courtown, the leeward fore-reef terrace at Albuquerque is
broader, extending for 1 to 1.6 km, and reaching depths greater than 30 m. It is an
12
extensive, gently dipping platform, descending at a low angle (4 to 7 degrees) to about 15
m and then gradually steeper to nearly 40 m, where the subvertical drop-off of the outer
slope begins. The bottom in the upper portions of the terrace is mostly covered by ripple
marked sand and rubble, although the calcareous hardground appears at certain locations
as elongated buttresses, about 1.5 m high, being thus sparcely overgrown by brown algae
(Stypopodium, Dictyota) and branching octocorals. Scleractinians are very scarce to
depths of about 12 to 15 m (living coral coverage: 5-20%), but their abundance and
species richness increase gradually with a simultaneous increase of the slope angle. At the
two localities visited, the outer margin of the terrace is marked respectively at -18 and -27
m by a subvertical escarpment, densely covered by plate-like and pagoda-like
scleractinians (Agaricia spp., Montastraea franksi), sponges and antipatharians, which
descend to nearly -35 m and give way to the accustomed sand step. Such a sand-covered
bench or step at 40-45 m depth was recorded on bathymetric profiles at other places of
the leeward outer margin of Albuquerque (Fig. 8), and can be distinguished on aerial
photographs as a lighter narrow band, outlining almost the entire outer slope around the
atoll shelf.
DISCUSSION
Rather than by its origin (e.g. Darwin, 1842), an atoll is defined by its geomorphic
features (Milliman, 1967, 1973; Geister, 1983). Hence, Albuquerque and Courtown may
be called atolls. When Milliman (1969) first described the gross morphology and
environmental features of the southwestern Caribbean atolls, he was impressed by their
close climatologic, oceanographic and geologic resemblance to many Pacific atolls:
surrounded by deep water, little seasonal change, appreciable windward fetch, and a
Millepora-Palythoa zone that emerges at low tide, resembling somehow the leeward
portions of the algal ridge found in Pacific reefs. Besides this, the atolls belonging to the
archipelago of San Andrés and Providencia are supposedly the only ones in the Caribbean
atolls with a volcanic basement (cf. Milliman and Supko, 1968; Geister, 1992).
The atolls of Albuquerque and Courtown share with nearby San Andrés Island and
other reef areas of the archipelago, the geological foundations upon which they rest and a
similar set of environmental conditions. Other Atlantic atolls, such as the ones found off
Belize and the Yucatan Peninsula (Lighthouse Reef, Glover's Reef, Turneffe Islands,
Chinchorro Bank, see Stoddart, 1962; James and Ginsburg, 1979; Jordan and Martin,
1987), in the Gulf of Mexico (Alacran Reef, Kornicker and Boyd., 1962) and the
Bahamas (Hogsty Reef, Milliman, 1967) show indeed some analogies with Albuquerque
and Courtown in their basic morphology, but they have different geological histories.
The presence of an extensive windward fore-reef terrace in Albuquerque and
Courtown is a characteristic common to most Caribbean atolls. As in the Belizean atolls,
the outer margin of the fore-reef terrace is defined by a sudden change of the slope angle
at about -20 to -25 m, where the nearly vertical cliff of the outer slope begins. The fore-
reef terrace or seaward bank is likely one of the essential morphological differences
between Caribbean and Pacific atolls. In the latter, the exposed reef margin margin is the
site of most active coral growth, leading to the development of characteristic shelf-edge
reefs.(cf. Wiens, 1962). The existence of a sandy step or bench at -35 to -45 m, that
outlines the outer slope of the atoll shelf, is also a common feature of the Belizean atolls
(cf. James and Ginsburg, 1979). This step, called by some authors the '-40 m Terrace’, is a
widespread characteristic of Caribbean reefs. It occurs also in the Bahamas (Zankl and
Schroeder, 1972), Jamaica (Goreau and Land, 1974), Curagao (Focke, 1978), San Andrés
(Geister, 1975), Providencia (Geister, 1992) and other Caribbean islands.
The present morphology of the outer margin in Caribbean reefs has been
interpreted in relation to the fluctuations of sea level in the last 80,000 years. As did James
and Ginsburg (1979) for the Belizean reefs, and Geister (1975, 1992) for the fore-reef
terraces of San Andrés and Providencia, respectively, we may assume that the outer
margin of Albuquerque and Courtown, indicated by the '-20 m Terrace’, corresponds to a
truncation of the former marginal reef area that occurred before the last interglacial
(Sangamon, about 125,000-80,000 years b.p.). In the period between Sangamon and
10,000 years b.p. sea level was not constantly low (about -120 m under present sea level).
At least three high stands of sea level took place during that time, reaching to nearly -25
to -40 m below present sea level (Bowen, 1988). The coincidence of the sandy bench in
present morphology at -35 to -40 m around both atolls, as well as the occurrence of a
deep intertidal notch at this level on the vertical cliff (at least at the visited locality in
Courtown), led us to explain this topography as a truncation of the emerging shelf margin
during a Pleistocene sea-level stand at about -40 m that may be regarded primarily as an
erosional feature. Unlike other Caribbean reefs, such as those off Belize (James and
Ginsburg, 1979) and Jamaica (Goreau and Land, 1974), where this feature is now
subdued by overgrowing modern facies, no significant accretion to the reef margin seems
to have occurred during the Holocene rise of the sea level either in Albuquerque or in
Courtown, or in the reefs surrounding San Andrés (cf. Geister, 1975) and Providencia (cf.
Geister, 1992), where a truncation of the outer margin at -35 to -40 m and an intertidal
notch are very distinctive.
The uppermost part of the reef front in both atolls shows a well developed spur-
and-groove system, similar to other reef complexes in the western Caribbean, such as
those off Belize, Yucatan, San Andrés and Providencia (cf. Stoddart, 1962; James and
Ginsburg, 1979; Jordan and Martin, 1987; Geister, 1975, 1992). In some localities, such
as the northeastern barrier of Courtown, where the relief between the spurs and grooves
often attains more than 3 m, and the grooves penetrate deeply into the reef flat, they are
apparently cut into Pleistocene rock, indicating an essentially erosional origin of this
system. It acts as an effective baffle for the immense energy expended by incoming surf
(Roberts, 1974; Geister, 1982). In other parts of the Caribbean, where the effective
windward fetch and the energy of the incoming surf are not as great, the spur-and-groove
system may owe much of its relief to differential rates of scleractinian growth (cf. Goreau,
1959). The presence of an extensive lagoonal terrace between the reef crest and the
lagoon basin on the windward side, as well as its abrupt transition into the lagoon in the
form of a 'sand-cliff, are also characteristics common to most oceanic reefs with a
considerable windward fetch, due to active movement of debris associated with the
14
prevailing northeasterly winds and waves. The presence of seagrasses on the lagoonal
terrace is conditioned by shelter created on the leeward side of the cays and islands.
The depth of the lagoon floor in Albuquerque and Courtown is not very different
from most Caribbean atolls, whose average lagoon depth ranges between 10 and 15 m
(Milliman, 1973). A singular feature is the existence of two well defined lagoon depth-
levels in Albuquerque. It is likely a consequence of the barrier effect of the 'Montastraea
spp.'-ribbon reef, which restricts leeward transport of bottom sediments to fill the lagoon
basin evenly. The occurrence of anastomosing and ribbon patch reefs covering unusually
large portions of the lagoon floor seems to be a rather common feature of oceanic reef
complexes in the Caribbean, such as Serrana Bank (Milliman 1969) and Alacran Reef
(Kornicker and Boyd, 1962). The NE portion of the lagoon bottom in Providencia Island
exhibits also several coalescing patch reefs (J.M. Diaz, J.A. Sanchez and S. Zea, pers.
obs., Sept. 1994). It seems likely that the greatest development of anastomosing patch
reefs is attained always on the windward side of the lagoon.
Contrasting with Pacific atolls, the absence or poor development of leeward
peripheral reefs is a characteristic common to most Caribbean atolls (Milliman, 1973).
However, Chinchorro Bank and some of the Belizean atolls exhibit a discontinuous
leeward reef crest which almost completely encloses the lagoon. Residence time of
lagoonal water may hence undergo a notable prolongation in these atolls. Coincidentally,
the abundance and development of lagoonal patch reefs in these atolls is apparently much
reduced in comparison to Albuquerque and Courtown (cf. Stoddart, 1962; Jordan and
Martin, 1987). It seems probable that the residence time of lagoonal water plays an
important part in the luxuriance and relative bottom coverage of patch reefs in Caribbean
atolls along with other physical factors, such as substrate availability and depth.
As stated by Milliman (1969), it seems probable that leeward peripheral reefs in
Albuquerque have originated from coalescing patch reefs. On the aerial photographs,
most peripheral reefs and the rubble zones surrounding them are arranged in a meander-
like fashion. Former ribbon and cellular reefs on the leeward lagoon margin have
apparently been damaged again and again by storms and hurricanes, leaving only the most
resistant frameworks of Acropora palmata and coralline algae, which built such peripheral
reefs. In Courtown, leeward peripheral reefs have developed only in the NW and SW
parts of the atoll, where the windward barrier reef bends southwestward at its northern
end and northwestward at its southern end. They are heavily exposed to colliding surf
from both the NW and the NE (or SW and NW) and are formed mainly by a framework
of coralline algae (Porolithon sp., Titanoderma spp., Lithophyllum sp.) comparable to
that of algal ridges. Although algal ridges had been thought characteristic of the Indo-
Pacific region until recently (Frost and Weiss, 1975), the southernmost leeward peripheral
reef in Courtown, with its emergent crest, represents in fact a true algal ridge, such as
those described recently elsewhere in the Caribbean (Glynn, 1973; Adey, 1975; Adey and
Burke, 1976). This feature was apparently overlooked by Milliman (1969), who refers to
it as a ‘small rocky spit, composed of massive coral debris’. Although not so well
developed, similar structures also have been recognized adjoining the NW end of the
barrier reef in nearby San Andrés by Geister (1975). Interesting discussions concerning
the existence and development of Caribbean algal ridges are found in Adey and Burke
(1976), Stoddart (1977) and Littler et al (1995).
At present, two cays exist in Albuquerque, both lying on the lagoonal terrace.
Their position, size, and shape have not changed significantly in the last 25 years, except
that North Cay has currently a more rounded shape than in the map of Milliman (1969)
and on the aerial photograph taken 1971. On the 1984-photograph it exhibits
approximately the current shape and size. The western and southern shores of this cay
have been dammed with piles of Strombus shells by the marines of the Colombian navy.
On the other hand, islands and cays in Courtown experienced remarkable changes in
number, size and shape since that time, and it seems likely that further changes are even
now taking place. Milliman (1969) mentioned four small cays, a sand spit and a rocky spit.
Sand Cay and East Cay lay close together on the lagoonal terrace and have currently
coalesced in an arrow-shaped island (about 800 m long), which seems to grow further to
the NW by accretion of sand and rubble (Plates 5 and 6). Of the formerly two cays sitting
on leeward peripheral reefs, Middle Cay was the only one visited by Milliman, who
noticed the presence of Yournefortia and Scaevola bushes and even some native
fishermen living on it. This cay might have disappeared between 1966 and 1971, since no
trace of it can be seen in the aerial photographs taken in August 1971. On the contrary,
West Cay (currently called Cayo Bolivar and serving as military post) and the sand spit
have experienced little change. The shallow bottom (1-2 m depth), where Middle Cay lies,
is currently covered with rubble and coral debris. It is not known if the.disappearance of
the cay was a slow erosional process that took place within five years or a rapid loss
produced by a forceful weather event. The latter seems less probable, since the only
hurricane recorded between 1966 and 1971 affecting this area, 'Irene' in 1971, had only
trivial consequences in nearby San Andres (IGAC, 1986).
Although a detailed checklist of scleractinians from Albuquerque and Courtown
has not yet been published, our survey indicates no noteworthy differences in species
composition and structure between the reefs of both atolls. It can be stated however that
the reefs in both atolls show a highly diverse fauna of about 40 species, not significantly
diverging from those known from neighbouring San Andrés and Providencia, where 44
and 43 species have been respectively recorded (Geister, 1975; 1992). The distribution
pattern of reef framework associations in both atolls, at least in shallow-water to about 15
m, is highly controlled by wave-energy and corresponds well to the ‘wave zones’ model
postulated by Geister (1977). With the exception of a ‘Porites zone’, each of the most
important reef framework associations recognized in the Caribbean Sea were found in
Albuquerque and Courtown. Only the names employed by Geister (1977) for his
'Melobesiae-zone' has been modified to designate the 'Coralline algae! unit (including the
algal ridges) in our maps.
In spite of a generally similar distribution pattern of reef framework associations,
there are some qualitative differences between Albuquerque and Courtown. Neither
'A.cervicornis' reefs nor a '‘coralline algae’ (or algal ridges) unit occur in Albuquerque.
Due to the interrupted windward reef crest in Courtown (ie.,discontinuity of the
‘Millepora-Palythoa” unit), medium-energy waves can penetrate in some places into
respective rear reef and lagoonal areas, leading to a better development of 'Diploria-
A.palmata" and 'A.cervicornis' reefs in this atoll. Contrary to Albuquerque, Courtown
lacks a gently dipping and extensive leeward terrace, which represents a highly abrasive
environment during storms and hurricans coming usually from the SW. This is seemingly
the main reason for a much reduced ‘scattered corals' unit and the lack of a ‘bare
calcareous hard bottom’ unit there.
Although detailed information about the current conditions of reef health in these
atolls will be presented and discussed elsewhere, some preliminary statements can be
made here. In the description of habitats presented above, we mentioned several signs that
are indicative of some degradation of the coral reef environment in both atolls.
Proliferation of algae overgrowing scleractinian colonies, low proportions of living coral
cover at several sites, abundance of heaps of skeletons of recently dead scleractinians (i.e.
Acropora spp.), as well as a noticeable depletion of commercial organisms, such as queen
conchs (Strombus gigas), lobsters (Panilurus spp.), snappers (Lutjanidae), groupers
(Serranidae) and turtles, are the most evident signs of degradation. Although no human
development exists in the atolls, they have been visited for many years by San Andrean
and Providencian natives for fish and turtles. In contrast to the condition in 1944, when
Fowler reported abundant fish and lobsters, by 1966 the populations of these resources
seemed to be low at Courtown, possibly a result of the increasingly fishing pressure
caused by overexploitation of Strombus, lobster and fish stocks at San Andrés (Wells,
1988). The health condition of reefs in these atolls is even at several sites not significantly
different from those around the densely populated San Andrés island (cf. Diaz ez. al.,
1995), indicating that, besides local human factors (i.e. sand mining, siltation, pollution)
and local natural agents (i.e. hurricanes), recent coral mortality is highly associated rather
to a generalized phenomenon of coral decline occurring in the Caribbean from beginning
of the 1980's (Hallock ez al., 1993; Ginsburg, 1994). Overfishing has also been recently
recognized as an indirect agent of coral mortality (Hughes, 1994).
ACKNOWLEDGMENTS
The authors express their gratitude to Luz S. Mejia, Guillermo Diaz (INVEMAR,
Santa Marta) and the crew of the R/V 'Ancon' for assistance in the field surveys. We
extend our appretiation to Dr. Jorn Geister (University of Bern, Switzerland) for his
helpful discussions and encouragement to make possible the flight over the atolls, as well
as friendly loan of the photos included in Plates 2, 4, 5 and 9. We thank Martha Prada for
her friendly hospitality at San Andrés. For their help in map digitizing and improvement of
computer drawings we are indebted to the students P. Sierra, J.A. Pulido and N. Ardila.
This study has been funded by the Instituto Colombiano de Ciencia y Tecnologia
(COLCIENCIAS, Grant No. 2105-09-023-93), the Instituto de Investigaciones Marinas y
Costeras (INVEMAR, Santa Marta) and the Universidad Nacional de Colombia.
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21
Table 1. Marine habitats (Map units) of Albuquerque and Courtown atolls with their
corresponding geomorphological units and absolute and relative area.
Habitat
unit
‘pioturbated
sand’
‘rubble with
algae’
‘sand and
rubble’
‘A.cervicornis’
‘scattered
corals’
‘rubble on hard
bottom’
‘Gorgonaceans
on hard bottom’
‘mixed corals’
‘Diploria-A.
palmata’
Geomorphol.
units
(depth)
Lagoon
(6-18m)
Lagoonal
terrace
(1-2m)
All zones
Lagoon
(3-5m)
Leeward
terrace
(15-30m)
Laggonal
terrace
(1-2m)
Fore-reef
terrace
(15-30m)
Fore-reef
and
Leeward
terraces
(25-37m)
Lagoon and
Lagoonal
terrace
(0.5-3 m)
Brief description
Calcareous sand (Halimeda, coral, shells)
with many burrows and mounds (Arenicola,
Callianasa).
Coral debris with rodoliths formed by
coralline algae mostly overgrown by brown
algae.
Bare coarse to medium sand with scattered
coral rubble and algal rodoliths.
Patch reefs dominated by thickets of
Acropora cervicornis, scattered coral heads
(Siderastrea, Montastraea) and plexaurid
octocorals.
Scattered massive and hemispheric
scleractinians (Siderastrea, Colpophyliia,
Diplona, Montastraea), gorgonaceans and
fleshy brown algae.
Rather barren bottom with coral debris
sometimes encrusted with coralline algae.
Rather flat bottom with luxuriant growth of
gorgonaceans (Pseudopterogorgia spp.,
Pterogorgia, Plexaurella, Eunicea,
Munceopsis, etc), scattered massive
scleractinians, many fleshy algae and large
sponges.
Diverse scleractinians (Montastraea franksi,
Diploria, Colpophyllia, Pontes, Mycetophyliia,
etc.) gorgonaceans, sponges and Halimeda.
Moderate wave exposed reefs dominated in
the shallow zones by thickets of Acropora
palmata, massive Diplona stngosa and
encrusting Porites astreoides.
Area (Ha)
Courtown Albuquerque
631.8 841.2
(12.8%) (11.7%)
127 122.4
(2.5) (1.7)
1270 237
(25.5) (33.2)
28.2 -
(0.6)
226.6 357.8
(4.6) (5)
'
361.3 260.2
(7.3) (3.6)
910 327.4
(10.3) (4.5)
134.1 180.2
(2.7) (2.5)
]
29.6 107.6
(0.6) (1.5)
22
Table 1. continued.
ae iad eos Brief description Area (Ha)
uni units
(depth) Courtown Albuquerque
‘ : Patchy seagrass meadows with Thalassia,
sealgiiass goonal Halodule and/or Syringodium growing on 35.5 3.8
aay sandy bottom. (0.7) (<0.1)
-om
‘ , Emerging sand and rubble accumulations,
land (cays) Fee mostly vegetated with shrubs (Scaevola, 9.2 78
eared Tournefortia), coconut palms or Ficus trees. (0.2) (0.1)
eewarl
peripheral
reefs
‘Millepora- Barrier reef Highly Wave-exposed reefs dominated by 195 195.4
; 0-3 m) Millepora complanata and Palythoa sp., (4) (2.7
Palythoa ( mostly accompained by crustose coralline 1)
algae.
‘Millepora- Barrier reef ear surf zone of the barrier reef. Millepora 107 59.2
pila 0-3 m) complanata, Portes ponites, P.astreoides (2.2) (0.8)
P.porites ( and Diploria strigosa. ; :
‘Montastraea Lagoon Ribbon and anastomosing patch reefs 325 1 304.1
‘ 5-15 m) dominated by massive Montastraea (6.5) (4.2)
Spp. ( annulans and M. faveolata, brown algae : :
(Lobophora-Dictyota) and some octocorals.
bare Fore-reet's War otioctigiemanidalytitcl) noggiiS.6 trafi224
calcareous hard terrace algae, scattered sea fans, brown algae. (8.4) (17.2)
bottom (3-15m) Heavily excavated by sheet-like sponges
(Cliona spp.).
‘outer slope’ Fore-reef Vertical to subvertical drop-off of the atoll 522.4 806.2
shelf. Sedimentary ramp or subvertical 10.4 11.2
and calcareous wall (covered or not with plate-like (10.4) (11.2)
Leeward scleractinians, sponges and antipatharians).
terraces
(>35m)
‘ : ’ Wave-exposed reefs, almost completely z
SEU I US cele Leeward covered by encrusting algae (Porolithon) - -
Peete building algal-ridge-like emerging crests. (0.5)
reets
(0-5m)
eee
4953.3 7172.5
a
23
BB RNK
EEA ACS
@)
es
CE HOS A
ASKS —S YA
Y SP
YS
LE
PAWN
ae
Lagoon with patch reefs
Windward barrier reef
ca Cays
oa Fore-reef terrace and outer slope
KA Leeward terrace and outer slope
¢| Lagoonal terrace
Leeward peripherical reefs
Sample sites
Figure 2, Geomorphological units and visited stations at Albuquerque Atoll. Straight lines
mark the location of the schematic profiles of Fig. 4.
24
81° 28'W
12° 28’N
Lagoon with patch reefs
| Ninaward barrier reef
ow Cays
[+ Fore reef terrace and outer slope
RY Leeward terrace and outer slope
0 500 1000 1500 m
Figure 3. Geomorphological units and visited stations at Courtown Atoll. Straight lines
mark the location of schematic profiles of Fig. 5.
A. OUTER LEEWARD TERRACE PR LAGOON LAGOONAL TERRACE] BR | fendez Ores
0 1 2 3
T4 15 16 7 km
PEN Fk Ce eRe od pe ta eer a pr ere erence REE
level
depth
= MO7TLLG aed SYP?
seattered corals
— mixed OTa
60 m L—
B. OUTER LEEWARD TERRACE lP.r| LAGOON Sty’ | LAGOONAL TERRACE B.R| F.R.TERRACE OUTER
(0) 1 2 3 4 5 6 7 8 9 10km
se siti
on ear ee ee
depth
bb with a
ip Montastraea spp.
gorgonian on hard bottom
re < QO
mixed corals
60 mL_—
Figure 4. West-East schematic profiles (straight lines in Fig. 2), showing the different
geomorphological and habitat units of Albuquerque Atoll. P.R- peripheral reefs, B.R-
barrier reef, F.R.- fore-reef.
25
26
§
3
>
© [ste |
ee | LAGOON LAGOONAL Terrace] 8.«| Seb
3)
i 2 4km
seq
depth
Millepora—Palythoa
Le coralline algae
rubble with algae
i gorgonians on.hard bottom ool of
Montastraea spp.
eee scattered corals
mixed corals
50 m
B. Lev [eri ¢ said GOON MMA Lee
0 Fr 19 aie 4 5 km
On == eye Oe SO ee ie Nim oe
depth
Millepora—Palythoa
= cervicornis
= algae or grass meadows
= coralline algae gorgonians on hard bottom
il Montastraea spp.
= scattered corals
mixed corals
50) tm ==
Figure 5. West-East schematic profiles (straight lines in Fig. 3), showing the different
geomorphological and habitat units of Courtown Atoll. P.R- peripheral reefs, B.R- barrier
reef.
77)
= Outer slope Z ward bottom with rubble
Fra Scattered corals Sea grasses
E24 Rubble with algae
—] Gorgonaceans on hard bottom Ej Diploria—A.palmata
12°07’N
=
lap)
w
2
2] Bare calcareous hard bottom Montastraea spp.
4 Millepora—Palythoa
| Bioturbated sediments
ai Millepora—P.porites EC] Sand and rubble
Figure 6. Distribution of bottom habitats and reef types at Albuquerque Atoll (for brief
description of map units see Table 1).
28
Diploria—A.palmata
Bioturbated sediments
= Millepora—Palythoa
A.cervicornis
fj Gorgonaceans on hard bottom
Eel Millepora—P.porites
(2) Bare calcareous hard bottom
Scattered corals
[_] Sand and rubble
E2 Montastraea spp.
@4 Coralline algae
[] Sea grasses
—] Outer slope
22 Mixed corals
Rubble with algae
Hard bottom with rubble
500 1000 1500 m
0
Figure 7. Distribution of bottom habitats and reef-types at Courtown Atoll (for brief
description of map units see Table 1).
Figure 8. Echosounder bathymetric profile of the leeward terrace and outer slope at
Albuquerque atoll. Note the presence of a truncation (sandy bench or step) at about -40 m
depth on the outer slope.
Plate 1. The spur-and-groove system of the windward barrier reef. The spurs are
overgrown on the top by Millepora complanata and by Porites spp. and crustose coralline
algae on the sides, whereas the narrow groove is filled with sand (Courtown, 22 May,
1994).
we
30
Plate 2. Oblique aerial view to the N, showing the buttress-groove system on the central
portion of the windward barrier reef at Courtown atoll (Sept. 29, 1994).
Plate 3. Rounded pinnacle (left) and narrow pillar formed by Millepora spp. at the SW
section of Courtown atoll, where the barrier reef becomes discontinuous (27 May, 1994).
31
Plate 4. Oblique aerial view to the W of Albuquerque atoll showing the two cays lying
close to the leeward margin of the lagoonal terrace (Sept. 29, 1994),
Plate 5. Oblique aerial view to the NW of Courtown atoll. The arrow-shaped island in the
center right is East Cay, which currently is connected with Sand Cay by a sand bar. Sand
Cay grows seemingly further to the NW by recent gradual addition of sand (Sept. 29,
1994).
32
Plate 6. East Cay, Courtown atoll, looking SE along the sand bar which at present
connects this cay with Sand Cay (21 May, 1994).
Plate 7. East shore of South Cay, Albuquerque atoll. Note the conspicuous band of
beachrock parallelling the shore line ( June 6, 1994)
33
Plate 8. Lagoonal patch reef in the upper depth level at Albuquerque atoll, made up
mostly by Montastraea annularis and M. faveolata (June 7, 1994).
Plate 9. Oblique aerial view to the SE of Albuquerque atoll. Note the two different hues
of the lagoon basin denoting the two depth-levels of the lagoon floor (Sept. 29, 1994).
)
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eu inet
ATOLL RESEARCH BULLETIN
NO. 436
CORAL FAUNA OF TAIPING ISLAND (ITU ABA ISLAND) IN THE SPRATLYS
OF THE SOUTH CHINA SEA
BY
CHANG-FENG DAI AND TUNG-YUNG FAN
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
APRIL 1996
115°E
Taiwan
¢ Pratas I.
Hainan
Paraceles I.
a o
rh)
%2¢ (a
‘ 1S)
15°N
Vietnam
SOUTH CHINA SEA
~
cA ~
7
2 : ~
, os XN
Taiping I. \ y Palawan
ping fe] :
114°21E
Taiping Island
E
Fig. 1. Locations of the survey sites (A-G) at Taiping Island
in the Spratlys of the South China Sea.
CORAL FAUNA OF TAIPING ISLAND (ITU ABA ISLAND) IN THE SPRATLYS
OF THE SOUTH CHINA SEA
BY
CHANG-FENG DAI AND TUNG-YUNG FAN
ABSTRACT
The coral fauna of the Taiping Island (Itu Aba Island) in the Spratlys of the South
China Sea was surveyed on April 19-23, 1994. A total of 163 species of scleractinians in
15 families and 56 genera; 15 species of alcyonaceans in three families and five genera; and
six species of gorgonaceans in four families and five genera were recorded. The coral
communities of the Taiping Island were dominated by scleractinian corals with high species
diversity and coral cover found on the lower reef flat at depths between 1 and 3 m.
Alcyonaceans and gorgonaceans are mainly distributed on the reef slopes at depths below
15 m. Wide reef flats and reef terraces exist on the east and west sides of the island
indicating that the reef development is better in these areas. Species diversity of coral
communities was the highest on the east side and the lowest on the west side of the island.
The depauperate coral fauna on the west side is possibly related to the strong SW monsoon
during summer and autumn. In comparison with other tropical coral reefs, species
diversity and abundance of coral communities of Taiping Island are relatively low. Dead
coral skeletons and debris were widely spread on the reefs below 3 m deep and only small
colonies were found. These facts indicate that coral communities of Taiping Island may
have been heavily damaged by natural catastrophes or artificial destruction during the last
decade. The possible destruction forces are typhoon disturbances and sea warming events.
INTRODUCTION
The South China Sea, situated between the Indian and Pacific Oceans, has an
historical importance in politics, economics, military affairs and transportation (Gomez,
1994). As the South China Sea is surrounded by continental Asia and many islands, it is
generally recognized as the major marginal sea in Asia. Major islands in the South China
Sea such as Tungsha Island (Pratas Island), Xisha Islands (Paracel Shoals) and Nansha
Islands (Spratly Islands) are reef islands. Most reef islands are atolls or emergent islands,
which are mainly composed of coral debris and sand. The emergent islands constitute only
a small portion of the reefs; the major parts are underwater reefs, shoals and banks.
Institute of Oceanography, National Taiwan University, P.O. Box 23-13, Taipei, Taiwan,
R.0.C.
Manuscript received 8 September 1995; revised 8 March 1996
The Spratly Islands, consisting of some 600 coral reefs and associated structures
scattered across an area north of Sabah and southern Palawan stretching for more than 500
km, are a group of atolls, islets, and sea mounts in the South China Sea. The structures
which protrude above the sea surface at high tide include at least 26 islands and seven
exposed rocks (McManus, 1992). Taiping Island, or Itu Aba Island, is one of the major
islands in the Spratly Islands.
The Indo-Pacific region, which includes the Spratlys, is characterized by a high
diversity of marine organisms. Among reef building corals, for example, the region in
which the Spratlys reside includes at least 70 genera (Veron, 1986, 1993). Inthe coral reef
ecosystem alone, more than 400 species of corals (Veron and Hodgson, 1989), 1500
species of reef fishes and 200 species of algae are found in this area (McManus, 1994). The
exact number of all marine species in the South China Sea is difficult to estimate given the
inadequate state of taxonomy, but the total number of species to be found at all depths in
the Spratlys certainly ranges to the tens of thousands (McManus, 1992).
The marine ecosystem of the South China Sea can be assumed to be dependent on
the Spratlys, at varying levels, for sources of larvae of renewable resources. Due to
prevailing monsoonal currents, the Spratly reefs may serve as sources of larvae that could
recruit to the disturbed coral reefs in the South China Sea (McManus, 1994). The semi-
enclosed nature of the South China Sea and hydrodynamic patterns prevailing in the area
could explain this linkage of coastal ecosystems in terms of nutrient level and fauna. It is
very likely that the Spratly Islands and similar groups of uninhabited reefs serve as a
mechanism for stabilizing the supply of young fish and invertebrates to these areas. This
becomes increasingly important wherein coastal populations of adult fish decline, as
appears to be the case in many coastal reefs of the Philippines and elsewhere. The dispersal
of larvae from the Spratlys possibly contribute to the coral reef fishery in the region. The
contribution of coral reef fishery to the national fish production of countries bordering the
South China Sea varies between 5-60% (McManus, 1994). Thus, the Spratlys could be
considered as a “saving bank” where commercially important fish and invertebrates are
saved from overharvest and supply a constant flow of larvae to areas of depletion.
Coral reefs are widely distributed in shallow water areas in the South China Sea.
The high spatial heterogeneity and productivity of coral reefs provide not only various
habitats for marine organisms but also feeding and nursery grounds for fishery resources
such as fish, shells, crustaceans and cephalopods. Flourishing coral reefs also constitute
beautiful underwater scenery that are valuable resources for the development of touristic
industry. As corals play a key role in marine ecosystems of the South China Sea, a better
understanding of the coral fauna in this area is necessary for conservation and management
of the marine resources in the future.
Several scientific expeditions in the South China Sea over the last 50 years have
provided oceanographic information and taxonomic listing of marine organisms, mainly
fishes. Although corals are widely distributed in the South China Sea, the coral fauna of
3
this area is poorly documented because of its remoteness and difficulty of access.
Bassett-Smith (1890) first described corals from Tizard Bank. Ma (1937) studied the
growth rates of scleractinian corals from Tungsha Island (Pratas Island). In recent years, a
few expeditions have been conducted to investigate the fauna and flora of the South China
Sea (Yang et al., 1975; Zou, 1978a, b; Fang et al., 1990). These studies have provided
valuable information for a preliminary understanding of the coral fauna of this area.
However, in comparison with the vast area of the South China Sea, these studies have only
covered a very restricted area. Studies on the coral fauna in other areas are thus necessary.
We sought to provide baseline information for resource conservation and
exploitation of Taiping Island (or Itu Aba Island). The objectives of this work were to
survey and to describe the distribution of coral reefs and reef topography, to provide an
inventory of coral species and their estimated relative abundance, and to identify special
coral biotopes.
STUDY SITE AND METHOD
Seven sites around Taiping Island (Fig. 1) were surveyed on April 19-23, 1994.
Taiping Island (10°23'N, 114°22'E), located on the northwest side of Tizard Bank, is one of
the major islands on the west side of the Spratly Islands (Nan-sha Islands). The island, with
an area of 0.49 km’, is about 1300 m long and 350 m wide (Fig. 3). The climate is tropical
oceanic. The average water temperature is about 28-29°C. The island is influenced by
seasonal monsoons. The northeast monsoon blows from October to March, the southwest
monsoon from May to October. The current flows southeast during the former and east or
north during the latter (UNEP/ITUCN, 1988).
Coral reefs were surveyed by snorkeling and scuba diving. Reef topography, coral
Species, community types and estimated coral cover were recorded. The relative
abundance of each coral species was estimated according to the number of colonies
encountered during each survey as common with more than 50 colonies, occasional with
about 10-50 colonies, or rare with less than 10 colonies. Underwater camera and video
were used to record photographs of coral colonies and reef topography. Coral species
were identified in the field. Whenever confronted with an uncertain species identification,
a piece of coral skeleton was detached and brought to the laboratory for further
identification. The identification of species was based on Veron and Pichon (1980, 1982),
Veron and Wallace (1984), Veron et al. (1977), Veron (1986), Dai (1989), Hoeksema and
Dai (1991), and Dai and Lin (1992).
RESULTS AND DISCUSSION
Description of Reef Topography and Coral Community
Site A is located on the south side of the island. The substrate of the upper reef flat
at 1-2 m depth is covered with sand and seagrasses. On the lower reef flat at 2-4 m depth,
there are abundant massive and stoutly branching colonies of Porites, Acropora and
Pocillopora spp. Below the reef flat at depths between 4 and 15 m, there is a steep slope;
only a few foliaceous Montipora and branching Acropora colonies were found on the
surface of the slope. At depths between 15 and 21 m, it is a reef terrace. The substrate is
flat and composed of coral debris with some ridges and grooves (Fig. 2a). The coral cover
is less than 5%; only a few small colonies are scattered on the substrate. The species
diversity was quite high, more than 67 species were recorded. The most abundant species
at this site is the octocorallian, /sis sp. (bamboo coral). They can form large colonies of 1
m long and in dense assemblages at some locations. Scleractinians found here are mainly
species of Montipora, Favia, Favites, Goniastrea and Cyphastrea. They typically exist as
small colonies with a diameter less than 10 cm. The widespread coral debris covering the
substrate was mainly Acropora and Pocillopora skeletons indicating that there were
flourishing branching coral communities in the past. The scarcity of coral species and
scattered small colonies indicate that the community might have been destroyed recently
and that recovery is slow.
Site B is located on the southeast of the island. Reef topography is similar to Site
A. There is a reef flat about 50 m wide at depths between 0 and 4 m. Living coral cover on
the reef flat exceeds 50%, but a trend of decrease toward the west is evident. Scleractinian
corals of about 120 species were found. Species commonly occurring on the reef flat were
stoutly branched colonies of Pocillopora damicornis, P. verrucosa, P. eydouxi, Acropora
monticulosa and A. gemmifera (Fig. 4). Colonies of A. digitifera, A, palmera, Favia
speciosa, Leptoria phrygia, Platygyra lamellina and the hydrocoral, Millepora
platyphylla were also commonly found on the reef flat. These species generally form large
colonies with diameters greater than 1 m. Corals existing on the flat are mainly massive,
encrusting and stoutly branched forms. The colony morphology of corals of this area
indicates that the reef flat is exposed to strong wave action. Below the reef flat on the
seaward side between 5 and 18 mis a steep slope on which coral cover was less than 5%;
only a few coral colonies were found to grow on the surface of the slope. A few solitary
corals of Fungia spp. and several large colonies of the blue coral, Heliopora coerulea,
were found on the sandy grooves. Below 18 m the bottom is sandy and no coral was
found.
Site C is situated on the west of the island. It is characterized by a wide reef flat
that extends westward to over 500 m from shore with depths about 3-8 m (Fig. 2b). On the
surface of the flat, there are low reef ridges alternating with shallow grooves running in the
NE-SW direction. Currents of this area are generally strong especially during flood and
ebb tides. This area is also exposed to strong waves during the summer monsoon. The
5
substrate on the upper reef flat was characterized by a dense seagrass bed. The lower reef
flat was covered with dead coral skeletons; some of them were clearly identifiable based on
skeletal features. Few small colonies were found and the coral cover was less than 2%.
These phenomena indicate that the coral communities might have been destroyed during
the past decade. Some small soft coral colonies such as Sarcophyton spp. and Lobophytum
spp. were scattered on the substrate (Fig. 5); few attained a diameter of 50 cm.
Site D is located on the northwest side of the island. The reef flat has a width about
100 m and stretches from 1 to 6 m deep (Fig. 2c). Coral communities on the reef flat can
be divided into two zones. In the upper zone between | and 3 m deep, coral cover is higher
than 50%. Species common in this zone are Favia, Favites, Goniastrea, Coeloseris
mayeri and Pavona spp. Some large colonies with diameters greater than 1 m were found.
In the lower zone between 3 and 6 m deep, coral diversity is low and coral cover is less than
10%. The reef surface is covered with dead coral skeletons and algae. Below 6 m, there is
a steep drop-off, descending at a nearly perpendicular angle to a depth about 60-80 m. On
the wall of this drop-off, there are colonies of Dendronephthya spp., Junceella fragilis and
Isis sp. Scleractinians were rare; only few small colonies of foliaceous corals were found
to grow on the slope. The coral cover is less than 5%. However, sponges, bryozoans and
other sessile invertebrates are abundant.
Site E is situated on the northeast side of the island. Reef topography and coral
fauna of this site are similar to those of Site D. On the upper zone of the reef flat, the coral
cover was higher than 50% and approximately 100 scleractinian species were found.
Among the most abundant species are Pocillopora verrucosa, P. eydouxi, Acropora
digitifera, Heliopora coerulea, and Millepora platyphylla (Fig. 6). Species of Montipora,
Porites, Favia, Favites and Goniastrea are also common in this zone; most of them are
massive, encrusting or stoutly branched forms, with colony sizes often less than 30 cm in
diameter. At the lower zone between 3 and 6 m deep, the substrate 1s covered mainly by
dead coral skeletons and green algae, Caulerpa spp. The coral cover is less than 5% in this
zone. There is a steep drop-off below 6 m; many large gorgonian and antipatharian
colonies were found overhanging on the slope. Sponges, bryozoans, crinoids and other
groups of marine invertebrates are abundant, which comprise a rich benthic fauna and
colorful scenery (Fig. 7). Below 35 m the bottom 1s sandy and no coral was found.
Site F is located on the east side of the island. The reef flat is wider in the north
where it extends seaward to approximately 500 m from shore but becomes narrower to the
south (Fig. 2d). Dense coral cover (>50%) and high species diversity were found on the
upper part of the reef flat at depths between 1 and 3 m. More than 100 scleractinian
Species were recorded, most of them were small colonies. Species commonly present in
this area are Pocillopora damicornis, P. verrucosa, Acropora digitifera, Cyphastrea
chalcidicum and Favites abdita. Coral cover and species diversity are low on the lower
part of the reef flat. Less than 5% of the substrate was covered by corals and only few
small colonies were found. Below 6 m there is a steep drop-off that extends to about 30 m
and reaches the sandy bottom. The most peculiar organisms on the surface of the slope are
6
many colorful soft corals, Dendronephthya spp. hanging on the wall. Other corals are rare
and scattered. Below 35 m the bottom is sandy and no corals were found.
Site G is located on a reef ridge on the southeast of the island. The reef ridge is
separated from the island by a trough approximately 20 m deep (Fig. 2e). The surface of
the ridge is smooth and about 7 m deep. More than 70 species of scleractinian corals were
found on the top of the ridge, mainly species of Acropora, Favia, Favites, Goniastrea, and
Fungia. The coral cover is about 30-40%. Many colonies of solitary corals such as
Fungia cyclolites, F. costulata, F. tenuis, F. fungites, F. scutaria and Herpolitha limax
were found on the sandy grooves. The edge of the reef ridge is about 8 m deep. Below 8
m there is a steep slope down to approximately 37 m. There are several 7ubastraea
micranthus colonies growing on the upper part of the slope. The lower part of the slope
between 20 and 37 m deep is covered by thick patches of Dendronephthya colonies (Fig.
8). These colorful soft corals, when fully extended, form a gorgeous underwater "flower
wall”. The slope reaches the sandy bottom at 37 m.
Coral Fauna
A total of 163 species in 15 families and 56 genera of scleractinians; 15 species in
three families and five genera of alcyonaceans; and six species in four families and five
genera of gorgonaceans were recorded during this survey (Table 1). The results showed
that coral communities of the Taiping Island are dominated by scleractinian corals with
high species diversity and abundant coral cover found on the reef flat between 1 and 3 m
deep. Alcyonaceans and gorgonaceans are relatively rare and their distributions are limited
to reef slopes at depths below 15 m. Although the coral fauna varied slightly among the
surveyed sites, species compositions of the coral communities are similar and can be
regarded as typical of tropical reef communities. The abundance of small coral colonies
indicates that coral communities are in their early stages of succession (Grigg, 1983). As
early succession communities generally have high species diversity (Connell, 1978), this
conditions may also relate to the high diversity of coral communities at Taiping Island.
In comparison with the known coral fauna of other reefs in the South China Sea,
the number of scleractinian species recorded during this study exceeds those of Tungsha
Island (Pratas Island, 101 species; Dai et al., 1995) and Xisha Islands (Paracel Shoals, 127
species; Zou and Chen, 1983). In general, the species composition of the coral fauna
among these islands is similar. Biogeographically, these coral fauna belong to the Indo-
Pacific province. Because Taiping Island is situated at a lower latitude and closer to the
area of highest coral diversity, it is natural that its coral fauna is more diverse than those of
other reefs in the South China Sea. According to the biogeographical location of Taiping
Island, this island is expected to have more than 70 genera and 400 species of scleractinians
(Veron, 1993). However, during our brief survey to the island, only 51 genera and 163
species were recorded (Table 1). Further intensive surveys of adjacent islands may reveal
more species.
7
The coral reef of Taiping Island is a typical oceanic reef. It has a wide, shallow reef
flat and a steep drop-off on the edge of the flat. The reef flat is a site of intensive coral
calcification that forms the reef framework. The substructure of this region is invariably
composed of large, massive, interlocking colonies of hermatypic corals cemented by
calcareous algae. The drop-off borders the reef framework and generally descends to
depths below 30 or 60 m. At the base of the drop-off there are abundant coral debris and
accumulation of sediment. These facts indicate that physical and biological destruction of
the reefs is relatively high and debris produced through these processes are transported to
a deeper zone at which accumulation occurs.
The development of reefs on the southwest and northeast sides of Taiping Island is
better than that of other areas. On both sides there are wide reef flats extending beyond
500 m from shore which basically conform to the shape of the island. Such a pattern of reef
development is likely related to the water flow of the reef as both sides are located in the
path of tidal current entering and leaving Tizard Bank. Reef growth is usually better where
there is strong water flow (Stoddart, 1969; Goreau and Goreau, 1973) because this flow
brings food and raw materials at the same time that it removes sediments and waste
products. In terms of species diversity, coral communities on the east, southeast and
northeast sides of the island are higher than in other areas. The depauperate coral fauna on
the west and southwest sides are possibly related to the strong SW monsoon during
summer and fall. Zou et al. (1978) reported that coral communities of Xisha Islands
(Paracel Shoals) were well developed on the northeast side and poorly developed on the
southwest side of the islands and that such distribution patterns are likely related to local
flow patterns. Due to the influence of the prevailing SW monsoon during summer, such
distribution patterns of coral communities are likely common in the South China Sea.
The tropical reef environment of Taiping Island implies that its coral fauna is rich
and the reef is highly developed. However, in comparison with other tropical Indo-Pacific
coral reefs, the species diversity and abundance of coral communities at Taiping Island are
relatively low. Dead coral skeletons were widely spread on the reef surface below 3 m and
only small coral colonies were found. These facts indicate that the coral communities of
Taiping Island have suffered severe damage during the last decade. The cause of such
extensive coral death is uncertain. Many natural and anthropogenic stresses on coral reefs
have been reported (see reviews by Brown and Howard, 1985; Grigg and Dollar, 1990).
According to the current status of the reef environment, the possible disturbances are likely
include artificial destructions, pollution, storms, predation of Acanthaster planci, and El
Nifio events.
Artificial destructions including blast fishing and underwater bombardment may
have caused heavy destruction in certain areas. The presence of idle troops at Taiping
Island is also of concern because they may engage in environmental damaging activities
such as shooting and fishing with explosives. Substantial damage may also come from
occasional parties of blast fishers and coral-smashing muroami fishers from the Philippines
and Vietnam (McManus, 1992).
The possibility of oil pollution is also of concern because the Spratlys lie near to
major shipping lines for oil and nuclear waste. Oil and nuclear waste could be released in
the event of a tanker accident in these reef-studded waters (McManus, 1992). However,
we found no substantial record or evidence of these pollutants.
The tropical position of Taiping Island places it within the area of frequent typhoon
disturbances. The typhoon-generated waves and storm surges may erode reef crest corals
and sediments down to about 20 m depth (Stoddart, 1985; Scoffin, 1993). The recognition
of past storm disturbances may rely on several features such as the deposits of coral debris,
the assemblages of corals and other reef biota, the reef framework structure, and the
existence of reef flat storm deposits (Stoddart, 1971; Scoffin, 1993). During this survey,
widespread coral debris were found to accumulate as talus at the foot of the fore-reef
slope, on submarine terraces and in grooves on the reef front. In addition, on the shallow
reef flat there are mainly massive, encrusting or stout branching corals that are basically
wave-resistant forms. These facts indicate that typhoon disturbances are possibly the
major destructive forces that have caused severe damage to the coral communities of
Taiping Island.
The population outbreak of the crown-of-thorn starfish, Acanthaster planci, has
been recognized as the most potent biotic disturbance affecting coral communities on many
Indo-Pacific reefs (Endean and Cameron, 1990). However, on reefs where marked
destruction of hard-coral cover was not apparent, A. planci was either not observed or
found at very low populations densities. Since we did not find any individual of A. planci
during this survey, it was unlikely that the crown-of-thorn starfish was the major
destructive force to the coral communities of Taiping Island.
Global sea warming associated with El Nifio events has caused widespread coral
bleaching in the Caribbean and the Pacific (Glynn, 1984, 1988; Williams and Bunkly-
Williams, 1990; Gleason, 1993). The ecological consequences of bleaching events include
widespread mortality with resultant decreases in coral cover, changes in species
composition, reduced growth rates and reproductive output of corals (Szmant and
Gassman, 1990; Gleason, 1993). Mortality rates in bleaching events have ranged from
zero (Hoeksema, 1991) to very severe (50-98%) as on the eastern Pacific during the
1982-83 El Nifio event (Glynn, 1988). This severe event also had other associated
secondary disturbances following coral mortality such as a subsequent increase in number
of grazers and bioerosion rates (Glynn, 1988). Whether the widespread mortality of corals
at Taiping Island is related to the El Niio-Southern Oscillation (ENSO) events need to be
studied. Analysis of the environmental record in coral skeletons and marine environmental
data are thus needed to answer this question.
In conclusion, the coral fauna of Taiping Island is dominated by scleractinian
corals, distributed mainly on the shallow reef flat at depths of 1-3 m on the east, south and
north sides of the island at which flourishing coral communities were found. Few
9
gorgonaceans and alcyonacean species were found mainly on deeper reef slopes. Coral
cover and species diversity of Taiping Island are relatively low in comparison with other
tropical Pacific coral reefs indicating that the coral communities of Taiping Island may
have been destroyed by artificial or natural disturbances. Since flourishing of coral
communities and reef-building activities are the basis of sustained development of this
island, we propose that reef conservation and protection are urgent and should be enforced
immediately by reducing artificial destruction and pollution to the reefs. In addition, the
changes of reef environment and biotic communities should be monitored. On a broader
scale, the Spratly Reefs, including Taiping Island, are ecologically important, with
abundant and relatively unexploited resources and where endangered species still abound.
The Spratlys may also serve as a pool of larvae for fishes and other marine organisms that
recruit to depleted fringing reefs and coastal habitats of the South China Sea. For these
reasons, it is worthwhile to conserve the ecosystem and genetic diversity of the Spratlys by
establishing a marine park in the Spratlys as proposed by McManus (1992).
ACKNOWLEDGEMENTS
We are grateful to Dr. L.-S. Fang, National Museum/Aquarium of Marine Biology
for his support and to Mr. D.-S. Chen for his assistance with field work. Special thanks are
due to the captain and crew of the Fishing Training Ship No. 2, Deep Sea Fishing Training
Center, Council of Agriculture. This study was supported by a grant from the Council of
Agriculture, Executive Yuan, R. O. C. (83-S.T.-2.15-F.-13).
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12
Table 1. Distribution and relative abundance of shallow water corals at seven study sites (A-F) of
Taiping Island. Relative abundance, +: rare, ++: occasionally, +++: common.
species / Site A B (C D E F G
SUBCLASS ZOANTHARIA
ORDER SCLERACTINIA
Family ASTROCOENIIDAE
Stylocoeniella armata + 3 + + +
S. guentheri + 3 + +
Family THAMNASTERIIDAE
Psammocora profundacella ar ase + + ++ ++ +
P. digitata 3° Te + +
P. contigua 2 35 25 + +
Family SIDERASTREIDAE
Pseudosiderastrea tayami = +
Coscinarea columna + + ~
C. exesa +
Famliy POCILLOPORIDAE
Pocillopora damicornis =r + ar a7 +++ ++
P. eydouxi 3 sete ++ + +
P. meandrina + 3 ae + 4 +
P. verrucosa oF states + ++ <FIaF +++ ae
P. woodjonesi +
Seriatopora caliendrum 4° +
S. hystrix + 35 + 36 + +
Stylophora pistillata 3 + + e 4
Palauastrea ramosa +
Family ACROPORIDAE
Acropora humilis absp 3 + +++ + a
A. gemmifera +++ + + + + +
A. monticulosa +4++ +++ + +
A. digitifera ar +++ ct ate +++ H+ +
A. robusta ++ + +f +
A. palmerae ++ + oF + +
A. nobilis ++ + ++ +
A. grandis + +
A. microphthalma + ain + +
A. aspera + +:
A. millepora +
A. tenuis a stot +
A. cytherea +
A. hyacinthus + + +
A. nana + +
A. cerealis et: +
A, nasuta +
A. valida qe + + ++ +
A. lutkeni 4 ae
A. divaricata a ie
A. florida +
A. sp 1 4
A. sp 2 su
Astreopora myriophthalma 4
A. listeri +
A. gracilis 4 +f + + + + +
Montipora monasteriata + ae + + +4 ae a
M. turgescens + 4 cfs a fh
M. undata +5 + + + a
M. verrucosa =H; qr + + ++ ++ a
M. danae 42
M. foveolata af
M. venosa + ++ + +
M. digitata oT +
M. grisea +
M. informis +P + + er iF +
M. foliosa + a
M. aequituberculata + + + +
Family AGARICIIDAE
Pavona clavus +
P. explanulata ++ + ++ a
P. varians + ++ ++ a
P. venosa + ++ + + +- ++ 4+
Gardineroseris planulata 4p + + +
Leptoseris mycetoseroides + + JL
L. explanata + +r
Coeloseris mayeri + + + + a
Pachyseris rugosa qe ++ ++ ++ - wee
P. speciosa + ++ + ated a dine
Family FUNGIIDAE
Fungia (Cycloseris) cyclolites cata ++
F. (C.) fragilis ns a
F. (C.) costulata + + + ats
F. (C/) tenuis + + a ffl
F. (C.) vaughani + ae =e
F. (Verrillofungia) repanda af + + + + +
F. (V.) concinna ae
F, (Danafungia) horrida 4
F’, (D.) scuposa i ay
F.. (Fungia) fungites ae + te spo
F. (Wellsofungia) granulosa a; te 1
F. (Pleuractis) gravis oe + + + a at
F. (P.) paumotensis at
F. (Lobactis) scutaria aF + + He nee i
species / Site A B (& D E F G
Ctenactis echinata + + ae ne
C. crassa x 24:
Herpolitha limax 3 2s + + +
Polyphyllia talpina + +
Sandalolitha robusta + + + - +.
Heliofungia actiniformis +
Family PORITIDAE
Alveopora verrilliana Efe a
A. spongiosa ss oy
Goniopora minor + 4:
G. columna ie aL
G. stuchburyi a +
Porites (Porites) solida ++ + + ++ + +
P. (P.) lichen + oF +
P. (P.) lobata + ++ + ++ ++ ne
P. (P.) lutea + ++ + ++ + 48
P. (P.) cylindrica + + + + zt
P. (P.) nigrescens + + a ae se
P. (P.) annae + +
P. (Synaraea) rus + ee + +
Family FAVIIDAE
Cyphastrea chalcidicum + +4++ + ++ +++ Fete
C. microphthalma + + + + +
C. serailia + ++ + + ++ +
Caulastrea furcata + -
Diploastrea heliopora + + + +
Echinopora lamellosa + + + + + as
E. gemmacea +
Favia favus sme + + oo = =
F. pallida + ++ + ++ ++ + a
F. rotumana +
F. speciosa + +++ a5 +++ ++ ++ +
F. stelligera + + - + +
F. laxa + a
Favites abdita + +++ + ++ ++ shekss ae
F. chinensis + + + +
F.. complanata +
F. flexuosa ++ + + ++ + +
F. russelli at ay be
F. pentagona + + + ++ + +
F. halicora +
Barabattoia amicorum + a
Montastrea valenciennesi or +
M. curta + ++ + + qa eet +
M. magnistellata + + x
Goniastrea edwardsi + + + + ra
G. aspera + +
G. pectinata ++ ++ + +
G. retiformis FF + + + =
Leptoria phrygia EF + ARP ote ser +
Platygyra pini staat + tte ay +
P. lamellina steht aaah ++ + ++
P. daedalea state + + ++ + +
P. sinensis ape +r atte tp +
Plesiastrea versipora ats ar +P
Leptastrea purpurea ats a +
L. pruinosa +f
L. transversa at +P +
Family OCULINIDAE
Galaxea fascicularis 2 ++ + 4 ++ ar “++
G. astreata +f alanis F + ++ ar +
Family MERULINIDAE
Merulina ampliata +P as af + +
Scapophyllia cylindrica ate +
Hydnophora exesa + ser + + ++ ++ ++
H. microconos qe ++ HP a
Family PECTINIIDAE
Echinophyllia aspera ct + + 4 + +
E. echinata + + oF +
Oxypora lacera + + ete +
O. glabra 4
Mycedium elephantotus +
Pectinia lactuca + 4p +
P. paeonia + anata ++ + +
Family MUSSIDAE
Blastomussa merleti +r
Cynarina lacrymalis ote
Scolymia cf. vitiensis + + +
Acanthastrea echinata + ++ + 4 =r + +
A. hillai + +
Lobophyllia hemprichii ale
L. corymbosa + +r +
Symphyllia recta ate a a
S. radians + +
S. agaricia + +
Family CARYOPHYLLIIDAE
Euphyllia (E.) glabrescens a0 at
Family DENDROPHYLLIIDAE
Turbinaria mesenterina oF
T. reniformis + +
Tubastraea aurea +
T. micranthus ae He rs
SUBCLASS OCTOCORALLIA
ORDER STOLONIFERA
Family TUBIPORIDAE
Tubipora musica 35 ++ 35 + ++ - ef
ORDER COENOTHECALIA
Family HELIOPORIDAE
Heliopora coerulea + sears ats + ao ++ ++
ORDER ALCYONARIA
Family Alcyoniidae
Sarcophyton ehrenbergi
S. trocheliophorum
S. glaucum
S. sp.
Lobophytum sarcophytoides
L. mirabile
Sinularia exilis ++
S. gibberosa *e
S. numerosa i
S. sp. 1
Sa Spe2 +
+++ 4+ 44+
++++ +4
+
+
+
Family Nephtheidae
Dendronephthya sp. 1 + + ++
D. sp. 2 = +
D. sp. 3 zt ae ef:
Family Xentidae
Xenia sp. + a
ORDER GORGONACEA
Family Isididae
Isis sp. qe ae i + + + -
Family Melithaeidae
Melithaea ochracea + + + ~
Family Subergorgidae
Subergorgia sp. ia + + ++
S. sp. + ~
Family Ellisellidae
Ellisella robusta + i
Junceella juncea + + +
CLASS HYDROZOA
ORDER MILLEPORINA
Family MILLEPORIDAE
Millepora platyphylla aF Tee I ++ +++ +++ +
M. tenera + + ee fe
M. intricata +++ ae mn
M. tuberosa +
Total No. of species 67 121 88 103 106 107 86
18
500 1000
b
(o)
30
a) 500 1000
E
< c d
2 2 :
c?)
a
40 40
0) 200 0) 500
wii Sea grass
xn Coral debris
««v Branching coral
a n Massive coral
= Soft coral
Gorgonians
cv Alcyonaceans
0 500 700
Distance from shore (m)
Fig. 2. Reef profiles and distribution of benthic organisms at the study sites.
a: Site A, b: Site C, c: Site D, d: Site F, e: Site G.
Fig. 3. Taiping Island, or Itu Aba Island, is a reef island about 1300 m long
and 350 m wide.
Fig. 4. Coral community on the reef flat at Site B is dominated by stoutly
branched colonies of Acropora spp.
19
20
Fig. 5. Some small soft coral colonies of Sinularia sp. scattered on the
substrate at Site C.
Yar OF : “od
ery Ae
Fig. 6. Coral community on the reef flat at Site E is dominated by stoutly
branched colonies such as Pocillopora eydouxi.
Fig. 8. Colonies of Dendronephthya sp. on the reef slope at Site G on the
southeast of the island.
21
ry ile
1 was
bh ala AN eodbiine at as waters on NT
ag eee” agi ; : : ) .
ATOLL RESEARCH BULLETIN
NO. 437
FIRST OBSERVATIONS ON THE FISH COMMUNITIES OF FRINGING
REEFS
IN THE REGION OF MAUMERE (FLORES - INDONESIA)
BY
MICHEL KULBICKI
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
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FIRST OBSERVATIONS ON THE FISH COMMUNITIES OF FRINGING REEFS
IN THE REGION OF MAUMERE (FLORES - INDONESIA).
BY,
MICHEL KULBICKI
ABSTRACT
Total fish counts were made along 6 transects on fringing reefs in the region of Maumere
(Flores - Indonesia). This represents the first description of fringing reef communities in this area
of the Pacific. A total of 255 species, distributed among 36 families, were recorded. The major
families were the Pomacentridae, Labridae, Serranidae, Acanthuridae and Chaetodontidae. The
number of species per station was high (96 species) compared to similar counts for fringing reefs
in New Caledonia. Density was 7.2 fish/m? and biomass was 187 g/m?. The average weight of
fish was low (21.7 g), with the Pomacentridae comprising 68% of the density. Large fish (over 40
cm) were scarce, possibly due to fishing pressure. The major contributors to the biomass were
Scaridae, Caesionidae, Acanthuridae and Pomacentridae. Carnivores had the highest number of
species followed by zooplanktivores and microalgae feeders. Most of the density consisted of
planktivores and microalgae feeders, whereas biomass was dominated by microalgae feeders,
zooplantivores and macroinvertebrate feeders. Small species with short life spans constituted
most of the density. The trophic structure and distribution of life-history strategies were very
similar to observations made on the fringing reefs of mainland New Caledonia, but were different
from those of fringing reefs of two isolated islands (Ouvea Atoll and Chesterfield Island). There
was a relationship between the number of dominant species and diversity. Structure of the
fringing reef fish communities was mainly linked to habitat type, in particular, terrestrial runoffs
could be a major factor.
INTRODUCTION
The reef fish fauna of Indonesia is one of the most diverse in the world, with over 2000
species. The Flores islands are at the eastern end of the Indonesian archipelago and are likely to
support a species diversity lower than the larger islands further west such as Java, Sumatra or
Borneo (species diversity decreases eastwards in the Pacific, and smaller islands tend to have
fewer species than large ones). Other than a recent checklist (Kuiter and Allen, unpublished),
very little is known of the reef fish communities of Flores. There is no account of the abundance,
biomass, size distribution, trophic structure and the life history strategies of the major reef fish
species in that region. The first objective of this article is to present a set of data relating to these
subjects that were obtained in the Maumere region in 1993.
The second objective of this article is to compare the species rich region of Flores with a
less diversified region (New Caledonia). Several questions come to mind when studying
ORSTOM - B.P. A5 Nouméa New Caledonia
Manuscript received 4 June 1995; revised 1 February 1996
i)
e702 i i ee A en Rg PO dS
TIMOR SEA
Figure | : map of the Maumere region. The 4 stations are indicated by a & on the map inset. The
numbers on the inset correspond to the transects.
3
communities found in a species rich area. For a given habitat, are there more species per unit area
than in a less diverse region with similar habitat? Are there more "dominant species " (species
making more than 2% of the density or the biomass) than in a less diverse region? Is the trophic
structure or the distribution of the life-history strategies different from those observed on fish
communities from a similar habitat but a different region? One of the major problems in
answering such questions is to develop comparable sets of data. In the present case, the data from
Flores were collected using the same methods as those used for a large set of data collected in
New Caledonia (Kulbicki et al, 1994a).
MATERIAL AND METHODS
During the Pre-Indo-Pacific Fish Conference in Maumere (November 1993), the author
had the opportunity to visit 4 fringing reefs and to perform 6 transects (Figure 1). The start of
each transect was chosen at random on the reefs and the transects were laid in the direction of the
slope. The transects were 50 m long. All fish, except the cryptic species (most Muraenidae,
Ophichtydae, Syngnathidae, Gobiidae, Blenntidae, Synodontidae, Scorpaenidae, Antenariidae)
and juveniles (newly recruited fish, usually less than 5 cm, but may be as small as 3 cm, i.e.
Chromis viridis), were counted. For each record, the species name, number of fish observed, size
of fish and distance of fish to the transect were noted. The size of the fish were noted in 1 cm
classes for fish less than 10 cm, in 2 cm classes for fish between 10 and 30 cm, in 5 cm classes
for fish between 30 and 50 cm and in 10 cm classes for fish more than 50 cm. The distances of
the fish to the transect were estimated in 1 m classes for fish less than 5 m from the transect, and
in 2 m classes for greater distances. Fish beyond 12 m from the transect were not counted. The
diver covered each transect only once. The average time per transect was 90 min. Densities were
calculated according to the method given by Burnham et al (1980) and Buckland et al. (1993).
Fish weights were estimated from length-weight equations (Kulbicki et al., 1994a). Biomasses
were estimated using these fish weights and the same method as for densities.
The diet of each fish species was either taken from the data used by Kulbicki et al.
(1994a) or from information in FISHBASE (Froese et al., 1992). Species with no direct
information available were assigned the same diet as the closest species within the same genus or
family for which dietary information was available. The food items are divided into 9 categories:
fish, macroinvertebrates, microinvertebrates, zooplankton, other plankton, macroalgae,
microalgae, coral, detritus. The diet of each species is distributed among these 9 food categories.
The percentage of each of these food items is taken into account when calculating the
contribution of a given species to a trophic category. For instance, if species A eats 50% fish and
50% microalgae, and if this species has a density of 0.1 fish/m?, the contribution of species A to
piscivory will be of 0.1 x 0.50 =0.05 fish /m?.
Each fish species was classified within one of the 6 life-history strategy classes defined in
table 1 (see Kulbicki 1992 for a discussion on this classification). For most species the
classification is given by Kulbicki et al. (1994a). For the remaining species, data from
FISHBASE (Froese et al., 1992) was used to assign the species to a given class. For a number of
species the information available was absent or too scant for a classification. In such a case, I
used the same classification as for the closest species within the genus or the family.
Each transect was divided into five sections of 10 m each. On each section the cover of
each of the substrate categories (see Kulbicki et al., 1993 for details of the method) given in
Table 2 was noted (the total for each section being 100%) for a 5 m wide strip. Algae and coral
cover were noted in the same manner.
RESULTS
The stations (Table 2) were between 3 and 7 m deep with a minimum depth of 1 m and a
maximum of 12 m. The substrate was characterised by a large proportion of rubble (debris, small
and large boulders) and a small coverage of sand, which was either muddy or coarse, no fine sand
being found. Rock formations were usually from eroded reefs and not of volcanic origin, as found
on land. Macroalgae were very scarce. Coral and alcyonarians were present in significant
amounts at only one station.
A total of 255 fish species, distributed among 36 families, were recorded (Appendix 1).
The families with more than 5 species accounted for 77 % of the total species seen (Table 3), and
only 6 families (Serranidae, Chaetodontidae, Pomacentridae, Labridae, Scaridae and
Acanthuridae) had more than 10 species. The number of species per transect (95.7 species),
density (7.1 fish /m?) and biomass (187 g/ m?) were high (Table 4), but average weights were
small (21.7 g) due to the dominance of Pomacentridae in the counts. Pomacentridae accounted for
16% of the diversity, 68% of the density and 9.5% of the biomass. One species, Pomacentrus
coelestis, formed 48.7% of the total density and four other Pomacentridae (Chromis amboinensis,
Chromis xanthura, Neopomacentrus azysron, Pomacentrus amboinensis) were among the 10
most important contributors to density. The other important species with respect to diversity and
density were in the Labridae, but no particular species in this family dominated in density. Most
species had a low number of individuals in the counts, even the planktivorous Labridae, which
are usually found in schools elsewhere in the Pacific. The major contributors to biomass were the
Scaridae and the Caesionidae. Most of the biomass for the Scaridae was made of juveniles, which
cannot be easily identified underwater, but two species, Scarus fasciatus and S.quoyi, formed
one-third of the Scaridae biomass. The Caesionidae, which are all schooling species, were
dominated by Pterocaesio tile and Pterocaesio chrysozona. One of the major contributors to
biomass was Pomacentrus coelestis, a very small fish (3 g average weight), but which was
present in extremely high densities.
The trophic structure can be considered in species numbers, density or biomass (Table 5).
Most species were carnivores (23.2% macrocarnivroes, 14.2% microcarnivores, 11.9 %
piscivores), zooplanktivores and microherbivores represented respectively 21.7 and 20.5% of the
species. Density was dominated by zooplanktivores (59.9%), followed by microherbivores
(17.2%). The other trophic categories had little importance with respect to density. Three
categories dominated biomass: microherbivores (34.9%), zooplanktivores (29.9%) and
macrocarnivores (19.3%). Coral and detritus feeders were low in all respects. The low numbers
for "other planktivores" are normal for reefs in the tropical Pacific. Macroherbivores were not an
important group. As is usually the case in the Pacific, this group exhibits little diversity and low
densities, but the large size of macroherbivores makes this category, at times, a significant
contributor to the biomass. In Flores, these fish were small in size, most likely because of fishing
pressure.
The distribution of the life-history strategies was dominated by the abundance of short-
lived species (classes | and 2) (Table 6). Short-lived species were also the most diverse; however,
species with an average life span (classes 3 and 4) were also represented by large number of
species. Biomass was evenly distributed between short and average life-span species.
There were major differences in the distribution of the life-history strategies among
trophic categories (Figure 2). In particular, zooplanktivores were essentially short-lived species
5
whereas, the long living species were mainly macrocarnivores and piscivores. Microherbivores
were split between many small, short-lived, species which dominated the density of this group,
and a few large longer-lived species (Scaridae, Acanthuridae), which made up most of the
biomass.
The average size of the commercially important species (essentially Serranidae,
Lethrinidae, Lutjanidae, Scaridae, Acanthuridae) indicates that there are very few large fish
(Appendix 1). In particular, not a single species with more than 10 individuals sighted, had an
average size > 40 cm. The size frequencies for the most abundant commercial species are given
on Figure 3. Most Serranidae were juveniles or small species. The Lethrinidae, Caesionidae and
Scaridae were small in size (sizes at least 30% less than average reproductive size). This could be
due to fishing pressure, but the high densities observed indicate that other factors could possibly
be involved.
DISCUSSION
The data set presented here are minimal and one should be cautious in generalizing these
results to a large area. In the absence of other comparable data from the Flores Islands or even
Indonesia, it is difficult to assess how representative are these results. In particular, it is
noteworthy that the stations were sampled in a leeward zone and that on the windward side of the
island the morphology of the reefs is very different, and it is likely that the reef fish communities
there would be different also. However, data from New Caledonia (Kulbicki et al. , 1994a)
indicate that even in a wide zone, reef fish communities from the same type of reef habitat share
much in common in species richness, density, biomass and structure.
The substrate found on the stations is typical of many fringing reefs in the region.
Indeed, in many cases terrestrial runoffs bring very fine sediment, and wave action induces the
formation of rubble and coarse sediment. The very low algae and coral cover is not unusual
either, especially in turbid areas.
It is difficult to compare the total number of species with other areas, because the
sampling effort was low. However, this number (255) is higher than observations made on
fringing reefs in Hawaii, 81 - 187 species (Hayes et al. , 1982) or French Polynesia, 80 species
(Galzin, 1985), which have been sampled much more thoroughly. These numbers are comparable
to the highest diversities found in New Caledonia, 168 - 252 species, but with a much larger
sampling effort (Kulbicki, 1992). The number of species /station is a better indicator, if the
stations are sampled in a similar manner. The only data (Table 7) that have been collected
according to the same methods are from Kulbicki et al. (1989, 1994a). The species richness
observed in Flores is higher than in any of the New Caledonian areas. It is estimated that there are
1140 reef and lagoon fish species in the Maumere area (Kuiter and Allen, unpublished), whereas
there are 940 species in the SW lagoon of New Caledonia (Rivaton et al. 1989), with 550 species
in the Chesterfield Islands (Kulbicki et al. , 1994b) and 630 in Ouvéa (Kulbicki et al, 1994a). The
families that are best represented in Flores exhibit considerable species diversity in most parts of
the tropical Pacific, but some families that contain many species elsewhere (Apogonidae,
Holocentridae, Scaridae, Acanthuridae) (Thresher, 1991) did not exhibit similar diversity in our
observations.
The densities observed in Flores are very high, especially for fringing reefs. Such
densities have not been recorded in this type of environment in the tropical Pacific (Kulbicki,
1991). However, most of this density is due to only one species, Pomacentrus coelestis, a
6
planktivore. Large densities of planktivores are common on reefs (Kulbicki et al. , 1994a), and
these species are usually short lived and experience large temporal variations. The other
components of the density in Flores are usually found on fringing reefs in the Pacific, in
particular, the Acanthuridae, Pomacentridae and small Labridae. This is confirmed by the few
published studies on fringing reefs in the Pacific that give a detailed account of the contribution
of the various species to density. In Hawaii (Hayes et al., 1982), the dominant species were two
Acanthuridae (A.nigrofuscus, Ctenochaetus striatus), followed by small Labridae (Thalassoma
duperrey, Gomphosus varius), the Pomacentridae being the third major component of the
Hawaiian reef communities. In French Polynesia, Galzin (1985) also found a majority of
Ctenochaetus striatus on the fringing reefs, the second most abundant species being another
herbivore, the Pomacentridae Stegastes nigricans. In New Caledonia, the composition of the
density varied from one zone to another. In Ouvéa (Kulbicki et al., 1994a) the most abundant fish
were Acanthurus nigrofuscus and Stegastes nigricans, followed by three planktivorous
Pomacentridae (Pomacentrus coelestis, Chromis chrysura, Chrysiptera cyanea). In the
Chesterfield islands (Kulbicki et al., 1989) the most abundant species were Mulloides
flavolineatus, juvenile Scaridae, Acanthurus nigrofuscus, Ctenochaetus striatus, three species of
Caesio and three Pomacentridae, all herbivores (Pomacentrus molluccensis, Stegastes nigricans,
Pomacentrus vaiuli). On the main island of New Caledonia (Kulbicki, unpubl.data), the major
contributor to density were planktivorous Caesionidae (Pterocaesio diagramma, P.tile), several
Pomacentridae (the two major ones being Chromis viridis and Dascyllus aruanus, which are
mainly planktivores), Acanthurus nigrofuscus, small Labridae (Thalassoma lunare, T.lutescens)
and juvenile Scaridae.
The biomass (187 g/m?) found in the Flores is high for fringing reefs. In Hawaii Brock et
al. (1979) found 106 g/m?, on the GBR (inshore reefs) Williams and Hatcher found 92 g/m?; the
results for New Caledonia are given in table 7. The distribution of the biomass can be compared
only to the studies from New Caledonia. There, the major contributors varied greatly from one
zone to another. In Ouvéa (Kulbicki et al., 1994a) the top three species in terms of biomass were
herbivores (Hipposcarus longiceps, Acanthurus blochii, Acanthurus xanthopterus); in the
Chesterfield Islands (Kulbicki et al., 1989) the top species were two herbivores (Kyphosus
vaigiensis, Naso unicornis) and a carnivore (Mulloides flavolineatus); and on the mainland the
main species were planktivores (Pterocaesio tile, P.diagramma) and herbivores (Acanthurus
nigrofuscus, Scaridae spp.). The similarity between Flores and New Caledonia is the presence of
Acanthuridae and Scaridae as major contributors to the biomass. The differences are in the
species involved, with larger species in New Caledonia than in the Flores Islands.
The comparison of some length frequencies (Figure 3) between Flores and New
Caledonia show that there is usually no difference in the size range. However, no small Siganus
doliatus were observed in Flores, which could be due to the season, small Siganus doliatus (less
than 15 cm) being found mainly during the dry season in New Caledonia. Monotaxis
grandocculis did not exceed 22 cm in Flores, whereas this species was found to reach 38 cm in
New Caledonia, with the largest sizes found on the barrier reef.
It is often assumed that the number of species contributing in an important manner (major
species; more than 2% in the present case) to the density or biomass decreases as diversity
increases (Richards, 1952 and Whittaker, 1964 in McIntosh, 1967; Spight, 1977; Wahington,
1984). The relationship is not clearcut, because it is often not specified which diversity is taken
into account: the observed diversity (number of species in the sample) or the potential diversity
(number of species in the region). The correlation between density and biomass for major species
exists both for the observed diversity and the potential diversity, but is not as good for the latter
gi
(Table 8 and Figure 4). This result suggests that highly diverse communities have lower numbers
of dominant species. In other words, one would expect the resources to be better shared and
utilised in these communities that in less diverse ones. Analysis of the trophic structure and of
distribution of the life-history strategies will in part answer this question.
It is difficult to compare the trophic structure found in Flores with most of the findings in
the literature, because the methods were very different from one study to another (Kulbicki,
1991). The data from New Caledonia were collected and analysed with the same methods used in
the present study and are, therefore, comparable (Figure 5). The distribution of species among
trophic categories (Figure 5a) is very similar in all 4 studies. However, Flores had more
zooplankton feeding species than the fringing reefs of New Caledonia. In density (Figure 5b) and
biomass (Figure 5c) the results from Flores and mainland New Caledonia are almost identical.
The latter two islands differ from Chesterfield and Ouvea, both of which are offshore islands, in
having larger numbers of zooplanktivores, lower abundances of microherbivores and carnivores,
and larger biomasses of zooplanktivores. This larger importance of zooplanktivores in the Flores
and mainland New Caledonia could be linked with high terrestrial runoffs (these islands have
similar land masses -10 000 and 20 000 km? - and average rainfall - 1500 to 2000 mm/ year).
There are also trends common to all four studies. In particular, coral feeders form 2-7% of the
species but account for very little in density or biomass. Detritus feeders and "other planktivores"
are never an important component of the trophic structure, whereas they form between 10 and
15% of the abundance or weight for the coastal (mangroves and estuaries) areas in New
Caledonia (Thollot, 1992). Fringing reefs and coastal areas are often adjacent in New Caledonia,
thus indicating that the trophic structure is greatly influenced by the substrate.
Very few studies on reef fishes have treated life-history strategies (Kulbicki, 1991;
Kulbicki et al., 1992, 1994a) or assimilated structures (ecological categories x size classes)
(Harmelin-Vivien, 1989). Kulbicki (1992), based on original data, compared life-history
strategies from several types of reefs across the Pacific using the same classification. The data of
the present study can be compared with data processed in the same way for fringing reefs in New
Caledonia (Figure 6).
The distribution of species among life-history strategies is almost identical for all reefs
(Figure 6a). This result could be expected from the findings of Kulbicki (1992), who
demonstrated that within the Western Pacific there were little differences in this structure at the
species level. Flores and mainland New Caledonia also have very similar structures in terms of
density and biomass (Figures 6b, c). In particular, they differ from the fringing reefs of the
islands of Ouvea and Chesterfield by having more class-1 species, which have the fastest
turnover. Conversely, Flores and mainland New Caledonia have a low proportion of biomass
represented by long living fishes (classes 5 and 6) which are important on the Ouvea and
Chesterfield islands. This suggests that in Flores the fish communities of the fringing reefs should
be more sensitive to short term variations than they would be on isolated islands such as Ouvea or
the Chesterfield. This is logical since most of these class 1 and 2 fish feed mainly on zooplankton
and microalgae, which are variable food sources, depending on primary production and mineral
inputs.
Our findings indicate, therefore, that the functioning of the fringing-reef fish community
of Flores is very similar to what is observed on mainland New Caledonia where ecological
conditions are similar. Conversely, fringing reef fish communities from isolated islands of New
Caledonia, despite their similar species composition, have different structures. Diversity alone
does not account for the major differences in the structure of these fish communities.
ACKNOWLEDGEMENTS
The author wishes to thank the following persons and organisations: Prof. Dr. Kasijan
Romimohtarto and the organizing committee of the Pre Indo-Pacific Fish Conference workshop
held in Maumere (November 20-25, 1993), R.Kuiter, Dr.G.Allen, G.Moutham, P.Dalzell and the
two anymous reviewers.
LITERATURE CITED
Brock R.E., Lewis C. et Wass R.C. 1979 Stability and structure of a fish community on a coral
patch reef - Marine Biology 54: 281-292
Buckland S.T., Anderson D.R., Burnham K.P., Laake J.L. 1993 Distance sampling, estimating
abundance of biological populations. Chapman & Hall London 446p.
Burnham K., Anderson D.R., Laake J.L. 1980 Estimation of density from line transect sampling
of biological populations. Wildlife Monographs 72: 202p.
Froese R., Palomares MLD, Pauly D. 1992 Draft user's manual of Fishbase software 7 -
International Center for Living Aquatic Resources Management- Manila Philippines 56 p.
Galzin R. 1985 Ecologie des poissons récifaux de Polynésie Francaise Thése Doctorat Université
de Montpellier: 195 p.
Harmelin-Vivien M. 1989 Reef fish community structure: an Indo-pacific comparison. in
Ecological studies - Vertebrates in complex tropical systems (Harmelin-Vivien M., Bourliére F.
eds) Springer Verlag N.Y. 69: 21-60
Hayes T., Hourigan T., Jazwinski S., Johnson S., Parrish J., Walsh D. 1982 The coastal resources,
fisheries and fishery ecology of Puako, West Hawaii - Hawaii Cooperative Fishery Research Unit
Technical Report 82-1: 159 + Annexes
Kulbicki M. 1991 Present knowledge of the structure of coral reef fish assemblages in the Pacific
- in Coastal resources and systems of the pacific basin: investigation and steps toward a
protective management - UNEP Regional Seas Report and Studies : 147: 31-53
Kulbicki M. 1992 Distribution of the major life-history strategies of coral reef fishes across the
Pacific. Proc. 7th Intern. Coral Reef Symp. - Guam 1992 : 918-929
Kulbicki M., Doherty P., Randall J.E., Bargibant G., Menou J-L., Mou-Tham G., Tirard P. 1989 -
La campagne Corail 1 du N.O. Coriolis aux iles Chesterfield (du 5 aoit - 4 sept. 1988) : données
préliminaire sur les peuplements ichtyologiques ORSTOM Nouméa. Rapp. Sci. Tech. Sci. Mer
Biol. Mar. 57 : 88 p.
Kulbicki M., Thollot P., Wantiez L. 1992 Life history strategies of fish assemblages from reef,
soft bottom and mangroves from New Caledonia. Seventh Intern Coral Reef Congress - Guam
June 1992 abstract
Kulbicki M., Dupont S., Dupouy C., Bargibant G., Hamel P., Menou J.L., Mou Tham G., Tirard
P. 1993 Caractéristiques physiques du lagon d'Ouvéa - in Evaluation des ressources en poissons
du lagon d'Ouvéa: 2éme partie: l'environnement physique: sédimentologie, substrat et courants -
Convention Sciences de la Mer ORSTOM Nouméa 10: 47-150
Kulbicki M., G. Bargibant, Menou J.L., Mou Tham G., P.Thollot, L. Wantiez, Williams J.T.
1994a Evaluations des ressources en poissons du lagon d'Ouvéa. in Evaluation des ressources en
poissons du lagon d'Ouvéa: 3éme partie: les poissons; Convention Sciences de la Mer ORSTOM
Nouméa 11: 448 p.
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Micronesica - 27 (1/2): 1-43
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Indonesian Journal
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Ecology 48 (3) : 392 - 404
Rivaton J., Fourmanoir P., Bourret P., Kulbicki M. 1989 - Catalogue des poissons de Nouvelle-
Calédonie. Catalogues Sciences de la Mer, ORSTOM Nouméa 2: 170 p.
Spight T.M. 1977 Diversity of shallow water gastropod communities on temperate and tropical
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des peuplements, relations avec les communautés ichtyologiques cotiéres. Ph.D. Thesis
University of Aix-Marseille II (France), 406 p.
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evolution, and possible implications - in The ecology of fishes on coral reefs (P.Sale ed.)
Academic Press Inc. New York 754 p.
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aquatic ecosystems - Water Research 18 (6): 653-694
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mid-shelf and outer shelf reefs of the Great Barrier Reef - Marine Ecology Progress Series 10:
239-250
10
Piscivores Macrocarnivores
Life-history strategy classes Life-history strategy classes
Microcarnivores Zooplanktivores
70 + Oce2p
eo Bic2B
50 +
40 +
x
30
20 +
10 +
(0)
1 2 3 4 5 6
Life-history strategy classes Life-history strategy classes
Microherbivores
Life-history strategy classes
Figure 2: distribution of trophic categories according to life-history strategies. D: density; B:
Biomass; Pi: piscivores; C1: macroinvertebrate feeders; C2: microinvertebrate feeders;
Zoo.: zooplanktivores; Mi.: microalgae feeders
Pterocaesio tile
number
16
Size (cm)
Scolopsis bilineatus
o
Ynumber
=
a
= Ss Sette Sea zz
Size (cm)
Ctenochaetus striatus
11
Monotaxis grandocculis
© Flores N=320
BI NC N=3825
Scarus fasclatus
O Flores N= 29 O Flores N= 23
BANC N= 1421 FNC N=295
Size (cm)
Figure 3: size distribution of the most abundant commercial species (NC:
Caledonia)
Y=10.13 -0.023X r1r?=0.60
12
i=
2 10
o
E =, 8
ore
06
ea
ow
iS
o
a
o Nn BR DD
100
Number of species in sample
200 300
Size (cm)
data for New
InY=6.95 -0.75InX 17=0.69
2.50 = In%B
2.00 5 |In%D
1.50
1.00
0.50
In % major species
0.00
6.00
400
7.50
In Number of potential species
6.50 7.00 8.00
Figure 4: correlation between number of species ("major species") contributing to more than 2%
of density (%D) or biomass (%B) and number of species in sample, or number of reef species
known in region. Data from Table 8. Note that for second figure a log scale is used.
WA
_] Flores Sp. ES Ouvea Sp. [J Chest. Sp. NC Sp.
% species number
a)
b)
”
7)
Oo
£
2
fe}
2 :
N .
=N %
EN =f i
oo 5 3) S o <= io} Ss
N 5 s O
fo)
C)
Figure 5: comparison of trophic structure (a: species, b: density, c: biomass) of fringing reefs
Flores with New Caledonia: Ouvéa (Kulbicki et al., 1994a), Chesterfield islands (Kulbicki et al.,
1989), main island (NC) (Kulbicki (1991). Pi: piscivores; Cl: macrocarnivores; C2:
microcarnivores; Zoo: zooplankton feeders; Other P.: other plankton feeders; MaH.: macroalgae
feeders; MiH.: microalgae feeders; Cor.: coral feeders; De.: detritus feeders
13
Flores S. ES Ouvea S. Chest. S. NC S.
Life-history strategy classes
a)
L] Flores D. ES OuveaD. [J Chest.D. N NCD.
70 7
60 |
> 50
@ 40
®
So 307
32
20
10
(0) ce ss CoE a
1 2 3 4 5 6
Life-history strategy classes
b)
Flores B. 3 OuveaB. [:] Chest. B. NC B.
Life-history strategy classes
c)
Figure 6: comparison of life-history strategy classes in Flores and New Caledonia. Key same as
Figure 5.
14
Table 1: definition of the 6 life-history strategy classes used for defining structure.
Life length can be considered as life expectancy (LSO after recruitment)
Class Size Reproduction Behavior Growth Mortality Life length
1 Small to Very early in life Most species Very fast High 0.5 to 3
medium __—~ Very high gonado-somatic school years
< 30cm index or reproductive Simple sexual
effort behavior
2 Small to 1-3 years old at first Often schools, Rapid initially Medium 3 to7 years
medium reproduction may be
< 30cm High gonado-somatic territorial
index Sexual
behavior may
be complex
3 Medium to 2-3 years old at first Often schools, Rapid initially Medium 3 to7 years
large reproduction seldom or through
> 30 cm High gonado-somatic territorial life
index Simple sexual
behavior
4 Small to Late in life Seldom Slow after Low 7 to 12
medium - Usually > 50 % maximum schools first years
< 30cm size at first reproduction Often reproduction
Medium gonado-somatic territorial _— initial growth
index often fast
5 Medium to Late in life Seldom Slow after Low 7-12 years
large Usually > 60% maximum schools first
> 30cm size at first reproduction Often reproduction
usually Low gonado-somatic index __ territorial Often rapid
>50cm initial growth
6 Large to Very late in life Almost never Veryslow Verylow > 12 years
very large Usually > 60% maximum schools especially
> 50cm size at first reproduction except for after
usually > 1m Often ovoviviparous reproduction reproduction
Low gonado-somatic index
15)
Table 2: composition of substrate. Depths in m. All other numbers are percentages.
STATION NUMBER
1 2 3 4 5 6 Total
SUBSTRATE
Sand - muddy 12 6 8}
Sand - fine
Sand - coarse WZ, 5) 5 17 11 8 10
Gravel and Debris 3 7 10 24 7 36 16
Small boulder 3 3 2 10 SY 16 14
Large boulder 23 3 4 7a) 18 34 22
Rock 47 41 71 28 3 5 33
Beachrock 8 8 3
TOTAL 100 100 100 100 100 100 100
ORGANISMS
Algae 5) 1
Coral 13 <1 <1 D,
Alcyonarians 15
DEPTH RANGE 3/9 2/9 2/4 2/10 1/12 7/9 1/12
Table 3: major fish families and their contribution to total diversity and comparison with New
Caledonia (NC)
Family Number of %total Species in | Family Number of %total Species in
species species common
with NC
Serranidae : Labridae :
Caesionidae 7 Da, 5 Scaridae 15 5.9 13
Mullidae 8 3a 7 Acanthuridae 16 6.3 £5
Chaetodontidae 15) D9) 13 Siganidae
Pomacanthidae 7 De 5 Balistidae
Pomacentridae 49 19.2 42 Total 197 77 170
16
Table 4: density (fish/m?) and biomass (g/m?) of the major families and species.
FAMILIES DENSITY BIOMASS
SERRANIDAE 0.099 6.36
Pseudanthias squamipinnis 0.047 0.17
Cephalopholis urodeta 0.013 0.99
Epinephelus fasciatus 0.010 1.10
LUTJANIDAE 0.021 3.95
Lutjanus decussatus 0.015 2S
LETHRINIDAE 0.025 Sle
Lethrinus harak 0.006 E92
Monotaxis grandocculis 0.012 1.85
NEMIPTERIDAE 0.040 4.28
Scolopsis bilineatus 0.021 1.67
MULLIDAE 0.042 9.48
Parupeneus indicus 0.003 4.53
Parupeneus trifasciatus 0.021 1.10
CHAETODONTIDAE 0.049 1.67
POMACANTHIDAE 0.044 29,
POMACENTRIDAE 4.954 18.4
Chromis amboinensis 0.163 0.64
Chromis xanthura 0.226 0.23
Neopomacentrus azysron 0.139 0.48
Pomacentrus amboinensis 0.074 0.31
Pomacentrus brachialis 0.103 0.63
Pomacentrus coelestis 3.468 10.4
LABRIDAE 0.374 7.86
Cirrhilabrus cyanopleura 0.027 0.11
Cirrilabrus sp. 0.027 0.06
Halichoeres melanurus 0.056 0.29
Novaculichthys taeniourus 0.004 1.09
Thalassoma amblycephalum 0.048 0.23
SCARIDAE 0.106 33:1
Scarus spp. juvenile 0.052 13.0
Scarus fasciatus 0.016 5.56
Scarus quoyi 0.014 6.94
ACANTHURIDAE 0.132 18.4
Acanthurus leucocheilus 0.033 2.50
Ctenochaetus striatus 0.059 5.60
Naso hexacanthus 0.008 2.14
SIGANIDAE 0.023 4.85
BALISTIDAE 0.065 4.84
TOTAL fas 187
Table 5 : trophic structure. All numbers are percentages.
CATEGORY DIVERSITY DENSITY BIOMASS
Piscivores 11.9 Mp 8.4
Macrocarnivores DD 43 19.3
Microcarnivores 14.2 6.5 3.8
Zooplanktivores Dey Se) 29.9
Other planktivores 0.1 0.1 0.1
Macroherbivores 2 0.1 0.8
Microherbivores 20.5 ee 34.9
Coral feeders Soe! 0.5 0.9
Detritus feeders 2.0 9.2 D0)
Table 6: distribution of the life-history strategies. All numbers are percentages. Classes refer to
the classification given in table 2.
LIFE-HISTORY STRATEGY DIVERSITY DENSITY BIOMASS
CLASS
1 10.0 61.6 8.2
D, 39.8 eS Sled!
3 16.1 5.8 36.4
4 Zk 3.8 3}
5) 10.0 1S 10.0
6 2.8 0.1 Med
Table 7: species richness (species /transect), density (fish/m?), biomass (g/m?) from fringing reefs
in New Caledonia (SW lagoon, Chesterfield and Ouvéa)(Kulbicki, 1991; Kulbicki et al., 1989,
1994a).
REGION SPECIES RICHNESS DENSITY BIOMASS
Chesterfield 64 QBS) 90/200
Ouvéa 85 2.4 340
SW Lagoon 55 2.2/5.8 61/155
18
Table 8: number of species (N) contributing to more than 2% of density or biomass for Flores and
other fringing reefs in the Pacific. Sampled species: number of species sampled. Potential
species: number of reef species known in the area; %N: percentage of N in the number of species
recorded during the survey.
1: Kulbicki unpublished; 2: Kulbicki et al., 1994a; 3: Kulbicki et al. 1989; 4: Galzin, 1985; Hayes
etaliy1982
Region N density %Ndensity Nbiomass %Nbiomass Sampled Potential Land are
species species (km?)
Flores 6 3) 10 3.9 255 1140 ~10 000
New Caledonia (1) 10 29 11 3.2 348 940 20 000
Ouvéa (2) 14 ies 8 4.3 152 630 130
Chesterfield (3) 14 10.8 10 7.8 130 550 10
Moorea (4) 6 I>) 80 630 130
Hawaii (5) 9 4.8 187 460 =500
19
Appendix 1: list of species observed. St: number of stations where species was observed; N: total
number of individuals seen; Sch.: average size of schools; Size: average size in cm
NAME
Taeniura lymma
Plotosus lineatus
Saurida gracilis
Synodus variegatus
Synodus dermatogennis
Synodus spp.
Sargocentron caudimaculatum
Aulostomus chinensis
Pterois antennata
Pterois volitans
Pseudanthias squamipinnis
Pseudanthias tuka
Anyperodon leucogrammicus
Cephalopholis argus
Cephalopholis cyanostigma
Cephalopholis leopardus
Cephalopholis microprion
Cephalopholis miniata
Cephalopholis sexmaculatus
Cephalopholis spiloparea
Cephalopholis urodeta
Epinephelus cyanopodus
Epinephelus fasciatus
Epinephelus hexagonatus
Epinephelus merra
Variola louti
Variola albomarginata
Pseudochromis exquisitus
Pseudochromis paccagnellae
Apogon fraenatus
Apogon nigrofasciatus
Cheilodipterus lineatus
Malacanthus latovittatus
Carangidae spp.
Caranx para
Caranx tille
Caranx spp.
Gnathanodon speciosus
Lutjanus decussatus
Lutjanus fulvus
Lutjanus rivulatus
Lutjanus vittus
Macolor niger
Caesio cuning
Caesio lunaris
Pterocaesio chrysozona
Caesio xanthonota
2
NPN NY HH KH KN DH WK KP KN KN KH RK KN WK K DN WK KK WWNnN DN WH OK NH RK We Ke eS
Nn
i=)
KS NOK DK KK WH eS
100
—
WD We WD HAwW NY NM VY
—
lon
Se UMW pe HP OW HN NY OY
18
232)
17.3
15.2
W267)
NAME
Pterocaesio diagramma
Pterocaesio teres
Pterocaesio tile
Plectorhinchus picus
Lethrinus olivaceus
Lethrinus harak
Lethrinus rubrioperculatus
Monotaxis grandoculis
Pentapodus caninus
Scolopsis affinis
Scolopsis bilineatus
Scolopsis lineatus
Scolopsis margaretifer
Mulloides flavolineatus
Parupeneus barberinus
Parupeneus bifasciatus
Parupeneus cyclostomus
Parupeneus indicus
Parupeneus macronema
Parupeneus trifasciatus
Upeneus tragula
Platax orbicularis
Chaetodon adiergastos
Chaetodon baronessa
Chaetodon citrinellus
Chaetodon kleinii
Chaetodon lineolatus
Chaetodon lunula
Chaetodon melannotus
Chaetodon ornatissimus
Chaetodon pelewensis
Chaetodon rafflesi
Chaetodon trifascialis
Chaetodon trifasciatus
Chaetodon vagabundus
Chaetodon xanthurus
Heniochus varius
Centropyge bicolor
Centropyge tibicen
Centropyge vrolicki
Genicanthus lamarcki
Pomacanthus imperator
Pomacanthus xanthomethopon
Pygoplites diacanthus
Abudefduf saxatilis
Acanthochromis polyacanthus
Amblyglyphidodon aureus
St
KBWNWNNK DHE NPE UWHWHE NEP HE NEP KP PEP SPE NHK DPEPRWRKE WHE DRE UEP NHK EP wD eG
= S| ne
i)
i)
—
ins
DNDN WOOWWN KH WD
NO
nA ©
20
NAME
Amblyglyphidodon curacao
Amblyglyphidodon leucogaster
Amphiprion clarkii
Amphiprion melanopus
Amphiprion perideraion
Chromis amboinensis
Chromis atripectoralis
Chromis atripes
Chromis viridis
Chromis chrysura
Chromis flavicauda
Chromis flavomaculata
Chromis margaritifer
Chromis retrofasciata
Chromis vanderbilti
Chromis spp.
Chromis xanthura
Chromis weberi
Chrysiptera rex
Chrysiptera rollandi
Chrysiptera talboti
Dascyllus aruanus
Dascyllus melanurus
Dascyllus reticulatus
Dascyllus trimaculatus
Discistodus melanotus
Neopomacentrus azysron
Neopomacentrus nemurus
Neopomacentrus violascens
Paraglyphidodo nigroris
Neoglyphidodon crossi
Plectroglyphidodon dicki
Plectroglyphidon lacrymatus
Pomacentrus alexanderae
Pomacentrus amboinensis
Pomacentrus bankanensis
Pomacentrus brachialis
Pomacentrus coelestis
Pomacentrus lepidogenys
Pomacentrus philippinus
Pomacentrus reidi
Pomacentrus simsiang
Pomacentrus sp.
Pomacentrus taeniometopon
Pomacentrus vaiuli
Cirrhitichtys falco
Paracirrhites forsteri
Sphyraena barracuda
Sphyraena japonica
Anampses caeruleopuncta
Bodianus mesothorax
2)
road
BWOrrP Ke HB PWWNYK WN DWW WH PP KP WWE NH WNKH KH UNDUADWK KEP NNN KH WHEN KEN HK HK NH WwW
NAME
Cheilinus celebicus
Cheilinus chlorourus
Cheilinus diagrammus
Cheilinus fasciatus
Cheilinus trilobatus
Choerodon anchorago
Cirrhilabrus exquisitus
Cirrhilabrus cyanopleura
Cirrhilabrus sp.
Coris gaimard
Coris schroederi
Diproctacanthus xanthurus
Epibulus insidiator
Gomphosus varius
Halichoeres argus
Halichoeres chrysus
Halichoeres hortulanus
Halichoeres melanurus
Halichoeres miniatus
Halichoeres prosopeion
Halichoeres podostigma
Halichoeres nebulosus
Halichoeres scapularis
Hemigymnus fasciatus
Hemigymnus melapterus
Hologymnosus annulatus
Hologymnosus doliatus
Labrichthys unilineatus
Labroides bicolor
Labroides dimidiatus
Macropharyngod meleagris
Macropharygodo ornatus
Novaculichthys taeniourus
Pseudocheilinu evanidus
Pseudocheilinu hexataenia
Pseudocheilinu octotaenia
Pseudodax mollucanus
Stethojulis bandanensis
Stetholulis interrupta
Stethojulis strigiventer
Stethojulis trilineata
Thalassoma amblycephalum
Thalassoma hardwicke
Thalassoma janseni
Thalassoma lunare
Scarus spp.
Cetoscarus bicolor
Scarus bleekeri
Scarus altipinnis
Scarus dimidiatus
Scarus flavipectoralis
~
oo
NK KE DK AADANNWNNRKP KF KF WK NWN DPRK KP PEN RP RP UN WNnNNK PK NWWN WD KN WHY
RWW We PNY W
oo
Wo
—_
(Gey ey ee ee
Nn
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10.4
— (oe)
—_— ~~) — 00
—
ee eo ee dO oe, oe, a
eet et
— ee Won fp ~TIN
1.3
NAME
Scarus fasciatus
Scarus forsteni
Scarus microrhinos
Scarus niger
Scarus oviceps
Scarus psittacus
Scarus quoyt
Scarus prosognathos
Scarus sordidus
Parapercis clathrata
Parapercis cylindrica
Parapercis multiplicata
Parapercis tetracantha
Ecsenius bandanus
Ecsenius bicolor
Ecsenius midas
Plagiotremus rhinorhynchos
Amblygobius rainfordi
Istigobius decoratus
Ptereleotris evides
Ptereleotris heteroptera
Valenciennea strigatus
Acanthurus mata
Acanthurus fowleri
Acanthurus dussumieri
Acanthurus nigricans
Acanthurus blochii
Acanthurus lineatus
Acanthurus nigrofuscus
Acanthurus leucocheilus
Acanthurus olivaceus
Acanthurus pyroferus
Ctenochaetus binotatus
Ctenochaetus striatus
Naso hexacanthus
Naso lituratus
Paracanthurus hepatus
Zebrasoma scopas
Siganus argenteus
Siganus canaliculatus
Siganus corallinus
Siganus doliatus
Siganus puellus
Siganus vulpinus
Zanclus cornutus
Rastrelliger kanagurta
Amanses scopas
Aluterus scriptus
Balistapus undulatus
Balistoides viridescens
Melichthys vidua
n
oo
BEA HBB BPWHAN WR DHY HE UANN HK HNN KKH KH eK Ke PN HEP Hanne aernweyr Oe WHE WDA |
23
NAME St
Odonus niger
Pervagor melanocephalus
Rhinecanthus verrucosus
Sufflamen bursa
Sufflamen chrysopterus
Arothron meleagris
Arothron nigropunctatus
WN NY Fe W
Canthigaster solandri
N
65
Sch.
16.2
aN
Size
13.7
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ATOLL RESEARCH BULLETIN
NO. 438
GROUPER DENSITY AND DIVERSITY AT TWO SITES IN THE REPUBLIC
OF MALDIVES
BY
ROBERT D. SLUKA AND NORMAN REICHENBACH
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
APRIL 1996
Tp)
a)
>
—
(am)
=]
<<
=
io GaN
noe otal?
GROUPER DENSITY AND DIVERSITY AT TWO SITES
IN THE REPUBLIC OF MALDIVES
BY
ROBERT D. SLUKA! AND NORM REICHENBACH?
ABSTRACT
The density and diversity of shallow-water groupers at Gaagandu, North Male Atoll and
Olhugiri, Thaa Atoll, Republic of Maldives was enumerated using visual transects. Four
different habitat types were surveyed: reef lagoon, reef crest, reef slope, and a well-
developed lagoonal reef. Twenty-two species in seven genera were recorded. Median
densities ranged from 7 to 23 grouper 240 m*. At Gaagandu Island, the reef slope was
repeatedly sampled using 20-m belt transects to estimate the efficiency and accuracy of
the sampling methodology. Fifteen transects were necessary to estimate the median
density of all species within 10% of the reference value and to develop a species list
containing 80% of the total number of species observed. The species observed varied
in their degree of site attachment. Those species which were most closely tied to their
habitat exhibited clumped spatial distributions while those species which ’roamed’ over
large areas had random spatial distributions. The number of transects necessary to
adequately characterize the median density of a species was related to the degree of
clumping in its spatial distribution.
INTRODUCTION
Groupers are an important fishery resource throughout the world and are important
predators in coral reef ecosystems. Approximately 30 grouper species occur in the
Republic of Maldives. Maldivians prefer to eat tuna and have not developed extensive
reef fish fisheries (Anderson et al. 1992). Total reef fish catch is approximately 3000
tons per year (Anderson et al. 1992). At present, we are aware of only two operations
exploiting groupers, one of which has had little effect on the grouper population (Sluka
unpublished data). A market has developed exporting groupers to other southeast Asian
countries and to supply many of the resorts located around North Male Atoll. It is
therefore likely that reef fish, especially groupers, will come under increasing
exploitation in the near future in the Republic of Maldives. Differences in catch
1 University of Miami, Department of Biology, P.O. Box 249118, Coral Gables, Florida
33124 USA
2 The Oceanographic Society of Maldives, Male, Republic of Maldives
Manuscript received 19 July 1994; revised 19 November 1994
2
composition during exploratory fishing were found between a southern atoll (Laamu) and
more northern atolls (Alifu and Shaviyani) (Anderson et al. 1992). Shepherd et al.
(1992) reported that the abundance and biomass of all species combined was lower on
reef flats that were mined than on unmined reef flats. However, the abundance and
biomass of fish on slopes adjacent to mined flats was greater than on slopes adjacent to
unmined flats. ‘Four grouper species were among the 20 fish which showed the most
dissimilarity between these slopes. Cephalopholis miniata and Variola louti had higher
biomass on slopes adjacent to mined flats, while Plectropomus pessuliferus and Gracila
albomarginata had higher biomass on slopes adjacent to unmined reef flats.
The difficulties in using visual survey methods such as transects has been reviewed by
other authors (De Martini and Roberts 1982; Bortone et al. 1986; Sanderson and
Solonsky 1986; Greene and Alevizon 1989). Various techniques for solving problems
such as transect width (Sale and Sharp 1983), transect length (Fowler 1987), duration of
the survey (St. John et al. 1990), and sample size (Sale and Douglas 1981) have been
developed. However, these studies usually involved sampling the whole community and
in many cases were specifically directed towards sampling patch reefs. Methodologies
for surveying serranids were examined by the Great Barrier Reef Marine Park Authority
(1979) and Craik (1981) for the Great Barrier Reef region. Groupers are relatively
sedentary and site attached. Survey methods must take into account their cryptic
behavior and the likelihood of having a patchy or clumped dispersion pattern. This
clumped dispersion could lead to misleading results if only a few samples are collected.
The number of samples necessary to accurately assess population density will depend on
the degree of clumping in their dispersion pattern.
The density and diversity of groupers was studied at two sites in the Republic of
Maldives and related to habitat preferences of the different species. The sample size
necessary to accurately estimate the density and diversity of groupers in a specific area
was examined using visual belt transects.
METHODS
Habitat characterization: The atolls were divided into three habitat zones: 1) lagoon, 2)
reef crest, and 3) reef slope. The habitat was characterized by recording the coverage
class of dominant substrate (sand, sand-mud, rubble, and hard reef) and lifeforms
(seagrass, algae, sponges, octocorals, and hard coral). Substrata and lifeform
information were collected by visually estimating the coverage in a belt of 1 m* quadrats.
Coverage was scored in the following categories: 1) < 10%, 2) 10 - 30%, 3) 30 - 70%,
and 4) > 70%. In order to convert to cm’ the midpoints of each coverage class were
summed for each quadrat and averaged.
Visual surveys: Prior to observation, the observer was trained to accurately estimate
length using models of fish with a known size-frequency distribution (Bell et al. 1985).
Visual surveys were conducted similarly to GBRMPA (1979). A 20-m transect was
placed in a haphazard fashion along a particular depth gradient (parallel to shore). An
3
area 6 m out from one side of the transect was intensively searched for all grouper
species and then the diver searched the other side in a similar fashion. The number and
size of all groupers observed were recorded. Groupers were placed in one of five size
categories: <5 cm, 5-15 cm, 15-25 cm, 25-35 cm, and >35 cm. The depth and time
of each survey were recorded. All of the habitat zones had similar sampling effort
except the reef slope at Gaagandu, which was more intensively surveyed. A distance of
approximately 300 m along the reef slope from 6 to 20 m depth was repeatedly sampled
in order to assess the number of transects necessary for reliable estimates of density and
diversity. Species identifications were made using Heemstra and Randall (1984), Randall
(1992), Randall and Heemstra (1991), and Allen and Steene (1987). When information
on species identification differed between sources, Randall and Heemstra (1991) was
used. Species presence/absense data was collected at Chicken Island, near Gaagandu,
for comparison.
Statistical analysis: Descriptive statistics, histograms, correlations and other calculations
were performed using Microsoft Excel® software. The frequency distributions of
numbers of groupers observed per transect (240 m”) exhibited various degrees of skewing
to the right (Figure la, b). Because of the skewed distributions, medians were
considered to characterize the densities better than means. Performance curves based on
cumulative medians and species-sample curves were used to determine the number of
transect replicates needed to obtain adequate density and diversity estimates for groupers
observed in the 48 transects from the slope area (Brower et al. 1990). Medians were
compared statistically using a Chi-square procedure (Zar 1984).
For species with median density estimates greater than zero, performance curves were
calculated. The performance curves calculated were considered to stabilize when all
subsequent cumulative medians fell between the 40th and 60th percentiles calculated from
the entire set of 48 transects. The least number of transects required to stabilize the
performance curve was considered the number of replicates required for a reliable density
estimate. This process was repeated 20 times, with the order of the 48 transects entering
the cumulative median calculation being randomized each time. Medians were then
calculated from the 20 estimates of replicates required to obtain a reliable density
estimate. The median estimates for required replicates were then correlated with the
species dispersion pattern using Morisita’s Index of Dispersion (I,) (Brower et al. 1990).
For the density and diversity of all species combined on the reef slope, performance
curves and species-sample curves were calculated. The number of replicates required
for a stable density estimate was determined in a fashion similar to that noted for
individual species except for the criteria used to determine performance curve stability.
Instead of using one level for determining stability, i.e. the 40th and 60th percentiles,
several levels were evaluated. These levels included 20% of the median (30 and 70
percentiles), 15% (35 and 65 percentiles), 10% (40 and 60 percentiles), and 5% (45 and
55 percentiles). If the median estimated from all 48 transects is considered to be the
reference median density, then these different levels for assessing performance curve
stability would indicate the accuracy of the median estimated from a given number of
transects. The number of replicates based upon the species-sample curves were also
4
48).
is urodeta (n
48) and (b)
Figure 1: Frequency distribution of number of grouper observed per transect for (a)
= Cephalopholis
Aethaloperca rogaa (n
a)
Asuenbel4
11S SMS IS
10
A. rogaa (#/transect)
b)
Prrrrrs
OOO II
RrEDLCEetirLeceeeenereLereeLieeLereeLee eles
PETRELELEMERILEL EERE LULLEL ELE LE LeELeerery
OOK OOK HO HHH III A MHI
ANIPIPIID IDI L ILI SL LL LIN IID LLLP LLDPE
DO SESASNAASSA XMAS
Asuenbel4
C. urodeta (#/transect)
5
assessed at various levels of percent of species observed. The levels included >70%,
>80%, =90%, and 100%. This process was repeated 20 times, randomizing the order
of the transects each time. Medians were then calculated from the 20 estimates of
replicates required for each level of percentage of species observed.
RESULTS
Habitat characterization: Gaagandu Island is located inside the main atoll ring of North
Male Atoll. The northern and western sides of the island are surrounded by a lagoon
approximately 50 m wide and approximately 2 m deep at high tide. The lagoon was
primarily rubble with very small areas of sand (Figure 2a). The rubble areas of the
lagoon were covered by turfing algae, had no soft coral or sponges, and very little hard
coral (Figure 2b). The reef crest consisted of large, eroded coral heads covered by algal
turf. The crest had only slightly higher hard coral cover than the lagoon and had very
low coverage of sponges and soft coral. From the crest, the reef sloped down steeply
to a sand flat at 30 m depth. The reef slope appeared to be divided into areas of high
vertical relief separated by ’landslides’ of rubble with sand. The reef slope had the
highest percentage cover of hard coral (approximately 30%) and low numbers of sponges
and soft coral. The southwestern portion of the island had a well-developed reef
consisting of a huge bed of Acropora sp. interspersed by massive coral colonies. This
reef is designated as reef 1 for further analyses. The depth ranged from 1-10 m at reef
1 and no substrate/lifeform data was taken at this site.
Olhugiri island is located on the northern edge of the outer ring of Thaa Atoll,
approximately 2.35 N latitude, 73.05 E longitude. The lagoon of the atoll stretches
approximately 50 m in each direction around the island. The northern side of the island
is open to the sea and has a reef crest which slopes steeply down to 50 m where the slope
becomes much gentler. The western portion of the island is lagoonal connecting to
another island without any deep passages. The inner side of the island has a reef crest
which slopes gently to about 10 m into a sand flat. The eastern portion of the island has
a channel about 10 m in depth which allows passage of water into the atoll. The outer
and inner reef crests were sampled for grouper density and diversity No quantitative
habitat data was collected at Olhugiri.
Density _and_ diversity of groupers: There was no correlation between any species
abundance, nor total abundance, with depth (minimum, maximum, or mean) or time of
day (p> 0.05) along the reef slope. There was a significant difference in the median
number of grouper observed per transect between sites (X? = 44.84, df = 4, p < 0.001,
Table 1). The slope at Gaagandu had the highest median density with 23 grouper
observed per transect. Excluding the slope data, the other sites had no significant
differences in the median number of grouper observed per transect (X? = 4.74, df = 3,
p > 0.05). The lagoon at Gaagandu had a median density of 5 and the lagoon at
Olhugiri 16. These two sites were not included in the density comparisons due to the
low sample size (2 and 4 transects, respectively).
6
Figure 2: Substrata (a) and Lifeform (b) coverage of the site at Gaagandu Island, North
Male Atoll. Open bars represent the slope area (n=100 1 m? quadrats), solid bars
represent the reef crest (n=100), and striped bars represent the reef lagoon (n=40). (a)
S = sand, RB = rubble, and HR = hard reef. (b) AT = algae, SP = sponge, SC =
octocoral, and HC = hard coral.
a)
Percent Cover
Substrata
b)
Percent Cover
\
\
N
N
\
\
\
\
\
NN:
NY
Lifeforms
qj
Table 1: Median, maximum, and minimum number of grouper observed per 240 m?
transect within each zone at the two island sites.
GAAGANDU OLHUGIRI
INNER OUTER
CREST SLOPE REEF 1 CREST CREST
MEDIAN 7 7a) 2 10 JUGS)
MAXIMUM 13 50 18 24 15
MINIMUM 3 11 4 3 7
The lagoon at Gaagandu was characterized by low diversity (4 species). There were 7
species observed on the reef crest, dominated by Cephalopholis argus and C. urodeta
(Table 2). Reef 1 was dominated by C. argus and Epinephelus merra. The slope had
the highest diversity with 17 species (also the largest sample size). Cephalopholis
miniata, C. leopardus, C. urodeta, E. spilotoceps, and C. argus dominated numerically
in decreasing order of importance. Along the slope the densities of ’roving’ species,
such as G. albomarginata, Variola louti, and Plectropomus spp., were probably
underestimated; these species were frequently observed swimming along the reef slope,
but outside transect boundaries. Overall, the species of Cephalopholis tended to
dominate numerically with many Epinephelus spp. being rarely observed. The
Epinephelus groupers commonly observed (E. spilotoceps, E. merra, and E. macrospilos)
were similarly colored, a white to cream background with brown spots or hexagonal
markings.
The inner reef crest of Olhugiri had 16 species present and the outer reef crest 15. The
dominant species on both reefs was C. argus, with a median number per transect of 7
inside and 6 outside (Table 2). C. leopardus and E. spilotoceps were the second most
abundant species on the inner crest, whereas C. urodeta was second most abundant on
the outer slope.
Length-frequency distribution: The majority of grouper observed in the lagoons at
Gaagandu and Olhugiri were small (5-15 cm Total Length (TL)). No groupers were
observed over 25 cm TL. The reef crest and slope had similar size - distributions (X?
= 7.07, df = 3, p > 0.05). The < 5 cm and 5-15 cm categories were combined due
to an expected value < 1 (Everitt 1992). The majority of grouper observed were 5-25
cm TL. On the slope the smaller grouper (5-15 cm) were dominated numerically by
Cephalopholis leopardus and C. urodeta. The largest fish observed on the slope (> 35
cm) were Anyperodon luecogrammicus, Aetheloperca rogaa, C. argus, Variola louti, E.
polyphekadian, and C. miniata. Fish observed were mostly less than 50 cm TL. Fish
greater than 50 cm were mostly V. louti and P. laevis. The larger grouper observed on
the reef crest (25-35 cm) were C. argus. Reef 1 had similar numbers of fish in the 5-15
cm, 15-25 cm, and 25-35 cm categories when compared to the other sites at Gaagandu
(X? = 0.43, df = 2, p > 0.05). Reef 1 had a larger percentage contribution of the >
8
Table 2: Median and maximum number of groupers observed per transect (median,
maximum) for Gaagandu slope (GS), Gaagandu crest (GC), Gaagandu lagoon (GL),
Gaagandu reef 1 (GR), Olhugiri inside crest (OI), Olhugiri outside crest (OO). The
minimum number observed per transect was zero except * = 3, + = 1, and # = 2.
% = species observed outside boundaries of transects
SPECIES GS GC” Gl. (GEKea Or OO
Number of transects 48 11 2 12 13 6
Aethaloperca rogaa 156 --- --- = 0,1 ee
Anyperodon luecogrammicus 1,4 0,2 --- 12 0,2 ---
Cephalopholis argus 3,11 Bhi vatgaas 5: naeOs te (OF
C. leopardus 4.5,16 --- --- 0,1 1,6 Ua
C. miniata 5: aes) ele -<aw yO ae
C. sexmaculata 0,2 = a ae aes ae
C. spiloparea 0,4 _ al ns is ae
C. urodeta De OMe LO pe aa (0) pane 72 S500
Epinephelus caeruleopunctatus 0,1 0,1 oo 0,1 0,1 Om
E. fasciatus = aes % ee es a
E. fuscoguttatus 0,1 ee — owe ie ae
E. macrospilos 0,1 | | I = --- ---
E. merra --- O'S ..2 B95; S18. 5169 (O22 =
E. ongus 0,1 --- O51 .051 oo ---
E. polyphekadian 0,2 --- --- 0,1 = 0,1
E. spilotoceps 314 03 -- 0,3 1,4 0.4
E. tauvina --- = =e ese 0,1 id
Gracila albomarginata 0,2 --- a = 0,1 ect
Plectropomus areolata --- --- = 0,2 0,1 0,1
P. laevis 0,1 --- --- --- 0,1 ---
P. pessuliferous = AY aes £5 en at
Variola louti 0,2 = —_ ? cic mr
9
35 cm category than the other sites at Gaagandu. These larger grouper were mainly C.
argus with a few A. luecogrammicus.
There was a significant difference in the length-frequency distributions of groupers on
the inner and outer crests at Olhugiri island (X? = 12.51, df = 4, p < 0.05). Many
small (< 5cm) C. leopardus were observed on the inner crest, whereas only 1 < 5 cm
C. urodeta was observed on the outer crest. There were more smaller (5-15 cm) grouper
and fewer larger (25-35 cm) grouper on the inner crest than would be expected if the two
size-frequency distributions were similar. Alternatively, there were fewer smaller (5-15
cm) grouper and more larger (25-35 cm) grouper on the outer crest than would be
expected.
Similarity index: The similarity in species composition was compared using Jaccard’s
coefficient, which is based on species presence/absence data (Table 3). The reef slope
at Gaagandu was most similar to reef 1 and Chicken Island (53%). The rest of the sites
at Gaagandu were less than 50% similar, with the lagoon the least similar to the reef
slope and reef 1. The Olhugiri reef crests were most similar to each other (82%).
Sample number: Seven species in the slope area had median densities greater than zero
(Table 2). The median number of transects necessary for a reliable density estimate
ranged from 2 to 16 (Table 4). The number of transects needed was related to the
degree the species exhibited a clumped distribution as indicated by their I, values (r =
0.73, p = 0.06). Two species, Anyperodon leucogrammicus and Aethelaperca rogaa,
had I, values which were not significant or nearly so; this indicated their dispersion
patterns were not significantly different from random.
These species required only a few transects to determine their density. In contrast,
the other 5 species showed various degrees of clumping and required more transects to
reliably estimate their densities (Table 4).
For all species combined, the number of transects needed for an accurate survey ranged
from 7 to 37 depending upon the level of accuracy desired for the median density and
the percent of the species observed (Figure 3). Increasing the number of transects from
7 to approximately 15 provided a large increase in the accuracy of the median density
estimate and percent of species observed. The accuracy of the estimate of median
density increased from 20% to approximately 10% of the reference median density, while
the percent of species observed increased from 70% to over 80%. Further increases in
the number of transects provided more moderate increases in the accuracy of the median
density estimate and percent of the species observed.
10
Table 3: Similarity matrix of Jaccard’s coefficient comparing the presence - absence of
species among survey sites.
SURVEY SITE 1 2 3 4 5 6 7 8
1. Gaagandu slope 1.00
2. Gaagandu crest 0.33 1.00
3. Gaagandu lagoon 0.11 0.38 1.00
4. Gaagandu Reef 1 0.53 0.46 0.14 1.00
5. Chicken Island 0.53 0.46 0.00 0.36 1.00
6. Olhugiri inside Os7e" 0°39" OC. O565 70°47) =£-00
crest
7. Olhugiri outside 0.68 0.29 0.06 0.59 0.60 0.82 1.00
crest
8. Olhugiri lagoon Only O38" Or53. O53) "Orte Ol25° Oro eee
Table 4: Morisita’s index of dispersion (I,) in relation to the median number of transects
necessary for a reliable density estimate for the 7 most common species of grouper
observed in the slope zone at Gaagandu Island, North Male Atoll. Chi-square test
Statistics and associated probability levels indicate whether or not the species’ dispersion
pattern was significantly different from a random distribution.
MEDIAN NO.
SPECIES TRANSECTS I, Ss P
Anyperodon Z 1.31 64.5 0.045
luecogrammicus
Aethaloperca rogaa 3 bel5 57.9 0.133
Cephalopholis u 1.44 119.0 < 0.001
spilotoceps
C. argus 11 1.47 111.4 < 0.001
C. miniata ihe. 1.34 136.9 < 0.001
C. leopardis 15) 1.38 141.7 < 0.001
C. urodeta 16 1.91 220.0 < 0.001
11
Figure 3: Number of transects needed to obtain a desired level of accuracy in
estimating the median number of groupers per unit area (dashed line) and percent of
all species observed (solid line) on the slope at Gaagandu (reference values are 23 for
median density and 17 for total number of species observed).
100 20
18
95
9 16
=
rye a
re) 14 s
$ 3
@ 85 12 <
C¢p) c
- $
S 10 6
8 80
o 8
75
6
70 4
0 5 10 15 20 25 30 35 40
Number of Transects
DISCUSSION
Habitat can be viewed on a number of different scales. The density and distribution of
groupers were related to within and among zone differences in habitat type. First, at the
macro-scale, there were clear differences in the density and diversity of groupers at
Gaagandu Island between the lagoon, crest, reef 1, and the slope. The slope had a
higher sampling effort so that rarer species were more likely to be observed. These
different zones vary in the amount of refuge available for groupers. The lagoon and crest
had little relief. The lagoon at Gaagandu has been mined for coral (M. Haleem pers.
com.). The lagoon at Olhugiri has not been mined extensively and still has large coral
heads. The density at the lagoon at Olhugiri exceeded all sites except the slope at
Gaagandu. This indicates that the lagoon at Gaagandu probably supported a much higher
density of groupers prior to mining. Reef 1 had high relief, but consisted mainly of
dense thickets of Acropora sp., which might have limited their use by certain species as
the interstices were probably too small for movement and hiding (the dense thickets most
likely inhibited the efforts of the surveyor as well). Harmelin-Vivien (1977) found that
spur and groove reefs at 6-18 meters depth had more species of fish and a higher biomass
than the deeper sloping platform.
12
Within the different zones the species were associated, to varying degrees, with specific
features. Some species had very little association with structural features of the zone such
as the species of Plectropomus, Variola louti, and Gracila albomarginata. These species
were observed to freely roam large areas generally > 15 meters deep. Variola louti was
not observed in caves or hiding in the Society Islands, but swam off the bottom (Randall
and Brock 1960). Gracila albomarginata was observed frequently in shallow water 5-10
m, however, Randall and Heemstra (1991) reported that this species was more abundant
in depths greater than 15 m. This species tended to swim along the slope and did not
appear to hide when frightened, but swam away, as is consistant with Randall and
Heemstra’s (1991) observations. Smith-Vaniz et al. (1988) also indicated that this
species was an active swimmer, not resting on the reef substratum. Plectropomus
areolata appeared more substrate attached; the younger ones were observed swimming
among the Acropora thickets on reef 1. The species of Plectropomus feed mainly on
fishes and tend to be less sedentary than most groupers (Randall and Hoese 1986).
Aethaloperca rogaa tended to be intermediate between these free-roaming species and the
more substrate attached species. Individuals tended to swim about freely, but would
often hide under coral heads and ledges when approached. They did not traverse long
distances as did the previously mentioned species, but would remain near a large coral
structure in the water column.
The reef slope contained areas with high coral relief, in between which occurred
*landslides’ of coral rubble and sand. Stoddart (1966) documented these same features
of Maldivian reefs. These rubble patches were frequently inhabited by small
Cephalopholis urodeta and, especially, Epinephelus spilotoceps. The latter species was
usually observed on the edge of these rubble patches near high coral relief rather than
out in the open. Epinephelus merra was abundant in the lagoons of the islands and at
reef 1. This species is similar to E. spilotoceps, being a demersal carnivore living under
ledges near the bottom of coral mounds and rubble (Hiatt and Strasburg 1960). E. merra
is typically found in shallow water on patch reefs in lagoons and bays (Heemstra and
Randall 1993). Many C. urodeta observed had a coloration with the posterior 1/3 to 1/2
of the fish black. Species descriptions of this fish indicate that the Indian Ocean variety
has only a dark caudal fin, but that in "dark habitats" in the Comoros Islands it was
uniformly brown (Randall and Heemstra 1991). Small specimens (< 10 cm) of C.
urodeta were observed in shallow water that appeared uniformly black or with a red head
region and black body posteriorly. Most of the individuals of this species conformed to
the species description in Randall and Heemstra (1991), however many followed this
pattern of more extensive black coloration on the posterior 1/3 to 1/2 of the body and the
soft dorsal and anal fins. Cephalopholis urodeta is strongly demersal and rarely ventures
away from shelter (Hiatt and Strasburg 1960). The most site attached of the slope
species was C. leopardus. It was always seen within patches of coral with closely set
*finger’ arrangements. When approached it would dart into the coral head. Anyperodon
luecogrammicus was often seen in pairs. Cephalopholis sexmaculata was observed only
in Caves as iS consistent with the observations of Randall and Ben-Tuvia (1983).
13
C. argus tended to have a higher density at shallower depths and dominated the diversity
on the reef crest. This species is one of the most common food fishes (Randall et al
1985), and is generally one of the most abundant piscivores at most locations thoughout
the Indo-Pacific (Randall and Ben-Tuvia 1983). It is more common on exposed rather
than protected reefs (Randall and Brock 1960) and prefers depths of 1-10 m (Heemstra
and Randall 1993). Shpigel and Fishelson (1989) found this species on the shallow reef
table and reef wall in the Gulf of Elat. Harmelin-Vivien (1977) observed C. argus at
depths of 6-18 m on spur and groove reefs and 18-25 m on the lower sloping platform
at Tulear. Cephalopholis miniata is abundant in deep lagoons and dominates coral knolls
that are isolated at depths of 17-33 m (Randall and Brock 1960). At one knoll off the
slope at Gaagandu at 30 m depth, this species was the most numerous of the groupers
observed. The grouper species observed on the reef crest and lagoon were in close
association with structural features such as overhangs and crevices (with the exception
of C. argus, which roamed about freely while darting into cover when approached). The
species observed in the lagoon were all similarly colored (brown spots or hexagons on
a light background) and tended to blend into the background of algal covered rubble.
Hiatt and Strasburg (1960) found E. macrospilos under large coral heads and rock ledges,
seldom far from cover. Our observations on this species in the lagoon at Gaagandu
support their findings. Epinephelus fasciatus was observed in the lagoon closely
associated with shelter. Fishelson (1977) observed this species near rocks in the lagoon
of the Gulf of Eilat (Aqaba) as well as in the fore reef.
The number of transects required to adequately characterize grouper density and diversity
is dependent upon the dispersion patterns and the desired levels of precision, accuracy,
and percent of the species observed in the community. A single visit to a reef is not
likely to record all species present, especially cryptic ones (Sale and Douglas 1981). An
analysis similar to that conducted here could be done on a preliminary set of transects
in order to determine the number of transects required. The number of transects should
be determined not only by the dispersion patterns of the species of interest, but also by
logistical constraints on effort. Collecting a large sample might increase accuracy
minimally and use time that could be applied to other sites (Bros and Cowell 1987). In
addition, if only species densities are required, the level of effort devoted to a particular
species could be tailored to the degree to which a species is clumped. Only a few
transects would be required to characterize the density of a randomly dispersed species,
while a species which is clumped would require more transects.
The groupers observed in this study appeared to have specific habitat requirements or
preferences. The dispersion of the groupers throughout the site is probably related to the
dispersion of their preferred habitat. Cephalopholis leopardus is strongly substrate
attached and its distribution was significantly clumped (Table 4). The clumped
distribution of the species is likely due to a clumped distribution of its preferred habitat.
Thirteen transects would be needed to adequately characterize the density of this species
whereas a species such as Aethaloperca rogaa which had a random distribution (Table
4), would need only 3 transects. A. rogaa is a species which is not strongly substrate
attached. However, our data on Anyperodon luecogrammicus does not follow this
pattern as it was randomly dispered, but appears to be strongly substrate attached. A
14
more detailed investigation of its habitat might reveal that it is a generalist in its
association with the substrate.
ACKNOWLEDGEMENTS
We gratefully acknowledge the help of Mohamed Haleem, Omar Maniku, Ahmed
Shakeel, and Steve Holloway. Without their contributions this research could not have
been accomplished. We also thank the men of Gaagandu and Olhugiri Islands for
helping with the research and providing a great living environment. The manuscript was
significantly improved by two anonymous reviewers. This project was sponsored by the
Oceanographic Society of Maldives.
LITERATURE CITED
Allen, G.R. and R.C. Steene. 1987. Reef fishes of the Indian Ocean. T.F.H.
Publications, New Jersey. 240 pp.
Anderson, R.C., Z. Waheed, M. Rasheed, and A. Arif. 1992. Reef fish resources survey
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ATOLL RESEARCH BULLETIN
NO. 439
EFFECT OF TYPHOONS ON THE LIZARD COMMUNITY OF A
SHELF ATOLL
BY
MICHAEL JAMES MCCOID
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
APRIL 1996
Cocos I.
~ 4 Km
is es ae |
Figure |. Map of the Mariana archipelago with the location of the study site.
EFFECT OF TYPHOONS ON THE LIZARD COMMUNITY OF A
SHELF ATOLL
BY
MICHAEL JAMES MCCOID!.2
ABSTRACT
Two major typhoons hit the southern Mariana Islands within an 11 month span and
provided a- unique, unplanned opportunity to investigate storm influences on the
herpetofauna of an atoll. Habitat specialists (Emoia atrocostata and Cryptoblepharus
poecilopleurus) endured the largest population declines because of habitat destruction. All
other species, particularly scincids, suffered less drastic population declines. The highest
population declines for all species occurred on the developed (resort) end of the island,
suggesting that removal and restructuring of typhoon-adapted vegetation allowed complete
overwash and local extirpations. Cumulative effects of typhoons suggest a resilience to
storm influences by atoll-dwelling reptiles.
The Mariana Islands comprise an archipelago of volcanic origin oriented north-south
roughly equidistant between New Guinea and Japan. There are 15 major islands, with the
northernmost (Farallon de Pajaros = Uracas) located at approximately 20°N, 145°E and the
southernmost (Guam) at 139N, 145°E (Fig. 1). Two km south of Guam, situated on the
southern portion of a coral lagoon, is Cocos (Dano) Island. This atoll has a maximum
elevation of 2 m and is approximately 100 m by 2 km. As of September 1992, forest
vegetation on Cocos Island was dominated by Cocos nucifera (Coconut Palm), Hernandia
sonora (no common name) and Casuarina equisetifolia (Australian Pine). Understory
vegetation in the forest was dominated by Carica papaya (Papaya) with ground cover
dominated by unidentified grasses and Ipomoea pes-caprae (Railroad Vine). Bordering the
surf / tidal splash zone on the windward side of the atoll were dense thickets of Pemphis
acidula (no common name). Vegetation on the developed (resort) northeastern 1/3 end of
the atoll was dominated by C. equisetifolia, C. nucifera, and ornamental trees and shrubs.
An historical record detailing vegetation was provided by Neubauer and Neubauer (1981).
The atoll has undergone substantial changes during the past half-century including the
development of a coconut plantation prior to WWII, construction of a U. S. military
installation (formerly occupying approximately 1/4 of the island), two resorts (occupying a
total of 1/2 the island; the present resort occupies only 1/3 the atoll), and at least three
typhoons since 1949 that overwashed the island (Neubauer and Neubauer, 1981; per.
obs.). Only about 1/3 of the island remains as atoll forest, albeit regenerated.
The climate of the southern Marianas is tropical with annual diurnal temperatures
ranging between 22° and 31°C (Anon., 1990). Rainfall is seasonal (Anon., 1990) with
1Division of Aquatic and Wildlife Resources, P. O. Box 2950, Agana, Guam 96910,
USA. ;
2Present Address: Caesar Kleberg Wildlife Research Institute, Texas A&M University,
Kingsville, Texas 78363, USA.
Manuscript received 19 July 1994; revised 25 April 1995
Z
most occurring between June and December. Typhoons in the western Pacific are common
and have been recorded on Guam in most months of the year (Myers, 1991). The typhoon
season on Guam is between June and December.
Information on the effects of typhoons on the fauna of atolls is minimal; Jackson (1967)
reported that insects and vertebrates persist despite catastrophic impacts and that lizards
"somehow have found sufficient protection". Damage to and recovery of vegetation is
better documented, with estimates of as long as ten years for a marked recovery (Wiens,
1962). In this unplanned study, I document changes in the herpetofauna of Cocos Island
after the cumulative effects of two major typhoons.
While typhoons are a yearly event in the Mariana Islands, two storms of severe
magnitude recently hit Guam within an 11 month span. Typhoon Russ hit Guam in
December 1990 and Typhoon Yuri in November 1991. Minimum sustained wind speeds
to attain classification as typhoons are the same as hurricanes (>74 MPH = 119 KPH) but
these storms had sustained wind speeds recorded at 175 MPH (281 KPH). Along
southeastern exposures (including Cocos Island), the direction that typhoons usually
approach Guam, maximum estimated wave heights were 9 m. Damage caused by high
winds and waves, in both typhoons, were substantial on Guam and catastrophic on Cocos
Island. Typhoon Russ totally overwashed the atoll, defoliated all broadleaf vegetation,
and downed an unknown, but large number of trees, particularly C. equisetifolia along the
windward side of the island. Typhoon Yuri inflicted similar damage including loss of a
substantial portion of the remaining C. equisetifolia on the windward side of the atoll. An
estimated 40-60% of C. equisetifolia on Cocos Island were cumulatively lost during the
typhoons. Another cumulative overt vegetation change observed was the virtual
elimination of the P. acidula thickets bordering the high energy zone on the windward side
of the atoll. An estimated 95% of the thickets were destroyed by Typhoon Yuri. Between
typhoons, dominant forest vegetation releafed, seeded, and a dense understory of C.
papaya and C. nucifera developed. Also during this period, the remaining P. acidula
thickets releafed. Due to Typhoon Yuri, the papaya and coconut palm understory was
destroyed and tremendous amounts of debris from the resort were strewn throughout the
forest. The dominant understory vegetation that emerged after the second storm was /.
pes-caprae.
The herpetofauna of the Mariana Islands has been characterized as depauperate (Rodda,
et al. 1991) consisting of a pre-western contact terrestrial reptile fauna of 13 species
(McCoid, 1993). Ten of these species occur on Cocos Island (Gehyra mutilata, G.
oceanica, Lepidodactylus lugubris, Perochirus ateles, Cryptoblepharus poecilopleurus,
Emoia cyanura, E. caeruleocauda, E. atrocostata, E. slevini, and Varanus indicus) and an
additional two species (Hemidactylus frenatus and Carlia cf. fusca), both introduced to the
Marianas (McCoid, 1993), are established on Cocos Island. At present, Cocos Island
possesses the most diverse reptile fauna (12 species) of any island in the Mariana
archipelago. Declines in the herpetofauna of the Mariana Islands were discussed by
Rodda, et al. (1991) but most species formerly found on Guam still occur on Cocos Island.
Although there are no native amphibians on the Mariana Islands, Bufo marinus is
established on Guam and Cocos Island.
The pre-typhoon reptile fauna on Cocos Island was not uniformly distributed in all
habitats. The gekkonids G. oceanica, H. frenatus, and P. ateles, were found in both
developed and forested areas (McCoid and Hensley, 1994), but differences in densities
between these habitats were not investigated. Gehyra mutilata and L. lugubris, however,
were far more common in the relatively undisturbed forested areas; | encountered only two
L. lugubris in the resort area during nocturnal surveys and no G. mutilata (G. Rodda, pers.
com., recorded these species in the forest). Scincids were also not evenly distributed in all
habitats. Carlia cf. fusca, perhaps introduced as recently as the late 1980's to Cocos Island
(T. Fritts, pers. com.) was found only at a boat landing and public park on the western end
8
of Cocos and at the resort on the eastern end on the island in early 1989. By mid-1990, the
species was observed in intervening habitats on Cocos Island. By early 1991 (see below),
the species was abundant in all areas. Cryptoblepharus poecilopleurus was most
conspicuous on the windward (east) side of the island where it commonly occurred on tree
trunks in C. equisetifolia groves (Hensley and McCoid, 1994). Generally, any tree with a
trunk diameter > 2.5 cm had at least one resident C. poecilopleurus. Emoia cyanura was
found in both resort and forest areas but was associated with sunlit, open habitat.
Expansive areas of dense undergrowth harbored few individuals. Emoia caeruleocauda
favored heavily shaded areas and was common in the forest and resort, but was
occasionally found in open areas. Emoia atrocostata was restricted to the high energy P.
acidula zone (total habitat 4 ha) on the windward side of Cocos Island. Emoia slevini only
occurred in forest (total habitat 9 ha) (McCoid, et al. 1995).
Qualitative surveys of the herpetofauna of Cocos Island were initiated in April 1989 and
initially consisted of nocturnal surveys for gekkonids, diurnal surveys for arboreal scincids
(both time-constrained surveys), and diurnal surveys for terrestrial scincids using
rubberbands. Time-constrained surveys (N = 5, between April 1989 and December 1991)
for C. poecilopleurus were limited from 15 to 30 min during which all lizards seen while
walking through C. equisetifolia groves were recorded. Time-constrained surveys for
gekkonids were conducted on the resort and lasted between 1.5 and 2 h during which all
lizards encountered along a predetermined route were either collected or recorded. In
September 1990, sticky traps (see Rodda, et al. 1993), which provide a mechanism to
estimate relative abundance, were first employed to sample terrestrial reptile faunas in
forested, resort, and beach areas of Cocos Island. Traps (10-80) were placed at five m
intervals and checked every 15 min at which time any lizards captured were removed.
Generally, sticky trapping spanned the time between 0700 and 1200 h. Rubber-banding
was only rarely employed after September 1990. After the December 1990 typhoon,
nocturnal surveys were discontinued (see below) and only arboreal diurnal and sticky
trapping survey techniques were used.
Pre-typhoon Russ herpetological surveys of gekkonids in the resort yielded a qualitative
estimated community structure (expressed as percentage of total number of lizards) of P.
ateles (4.5 %), G. oceanica ( 6.2 %), L. lugubris (0.6 %), and H. frenatus (88.8%) (N =
315 lizards in 30 person-hours survey effort). Unfortunately, the survey route for
gekkonids was completely destroyed by the cumulative effects of both typhoons. This was
exacerbated by the clean-up efforts of the resort corporation in which remaining debris was
removed. Thus, no comparable post-typhoon data could be generated.
Surveys immediately after Typhoon Yuri yielded no lizards of any species on the
approximately 1/3 of the island occupied by the resort. This portion of the island was
subjected to the most intense vegetation / structural loss from typhoons. Although
gekkonids were common in the resort prior to the typhoons, population densities of
gekkonids in the relatively unsurveyed forest sections of Cocos Island are unknown; I can
only assume that a sizable fraction of the gekkonids on Cocos Island were lost because of
typhoons. Post-Typhoon Yuri diurnal surveys in forest areas targeting gekkonids revealed
the persistence of all previously recorded species on Cocos Island.
Pre-typhoon sticky trapping surveys for E. atrocostata yielded a Catch-Per-Unit-Effort
(CPUE) of 0.304 lizards/trap hr (N = 51 lizards, trap hrs = 168). Trap-hours are defined
as one trap set for one hr = one trap hr. CPUE's are the number of lizards captured/trap hr.
Post-typhoon surveys yielded a CPUE of 0.022 (N = 2 lizards, trap hrs = 90). This is a
decline of an order of magnitude in catch rates and suggests that the population on Cocos
Island declined by over 90% due to cumulative typhoon effects.
The remaining Emoia species (cyanura, caeruleocauda, and slevini) and C. cf. fusca can
be discussed as a group as no changes in ranking of species collected (see below) in the
forest area were noted after or between typhoons. These four species were initially
4
sampled in forest using rubber-banding in early 1989 through late 1990 and sticky trapping
in September 1990. Initial levels of efforts were low (total trap hrs = 22) or not
quantifiable (rubber-banding). Numbers of lizards collected, ranked in terms of most to
least abundant, indicated that C. cf. fusca was the most common followed by E.
caeruleocauda, E. cyanura, and E. slevini. All sticky trapping surveys in the forest after
December 1990 (N = 5) were conducted along the same transects and yielded the same
ranking in abundance as above. Trapping (N = 1400 trap hrs) was conducted in January,
June, October, and December 1991, and September 1992. Two surveys (January 1991
and December 1991) were conducted within two weeks after typhoons. Percentage
composition for each of the species (grand total = 365 skinks) in the five forest surveys
ranged between 57.6 and 68.9 for C. cf. fusca, 20.7 and 30.3 for E. caeruleocauda, 2.6
and 12.9 for E. cyanura, and 0.0 and 2.6 for E. slevini. Changes in percentage
compositions between surveys were tested using a R X C test of independence with a
William's correction and were not significantly different (X7cale,12,.05 = 9.197). This
suggests that responses of individual species to typhoon effects were not statistically
different. Similarly, CPUE's for all surveys were within the same order of magnitude
(range 0.171 - 0.475) indicating that the cumulative effects of the typhoons did not
dramatically decrease catch-rates of forest-dwelling scincids. Since at least 1/3 of the island
was devoid of any lizards after Typhoon Yuri (see above), it is safe to assume that total
population declines were greater for E. cyanura, E. caeruleocauda, and C. cf. fusca than
for E. slevini, which occurred only in forest.
Numbers of C. poecilopleurus were gauged by sightings per min (range 0.33 - 1.1).
These sighting data, including both pre- and post-typhoon observations, are within the
same order of magnitude suggesting that typhoon effects were minimal on survivorship of
C. poecilopleurus. importantly though, post-typhoon observations were made on existing
trees and since sighting rates after typhoons did not increase on these trees, perhaps
indicating emigration of surviving lizards from felled trees to existing trees, it is assumed
that if a tree was lost during a typhoon, the resident lizards were also lost.
The ability of a herpetofauna to persist on an atoll after substantial environmental
perturbations are also highlighted by observations on two species not directly surveyed in
this report. Varanus indicus, although found on Guam, was probably introduced to Cocos
Island in the late 1980's (pers. obs.) and managed to persist through two major typhoons.
By December 1991, in addition to a number (3 - 5) of 200 to 450 mm snout-vent length
(SVL) lizards, a small (ca. 100 mm SVL) individual had been observed on Cocos Island.
These observations suggest that successful reproduction had occurred and monitor lizards
had survived the typhoons. Bufo marinus was probably introduced to Cocos Island in
1989 and successful reproduction (large numbers of tadpoles in rain pools) was observed
in September 1989. In September 1992, after both typhoons, two adult (ca. 830 mm SVL)
B. marinus were observed in a freshwater pool.
Observations of the herpetofauna on Cocos Island after typhoons suggest a resilience to
environmental perturbations. Terrestrial forest-dwelling scincid populations appeared to
persist relatively unscathed despite substantial typhoon impacts. Habitat specialists (E.
atrocostata and C. poecilopleurus) were more susceptible to population declines due to
habitat destruction. All gekkonid species also persisted after the substantial effects of the
typhoons. Besides C. poecilopleurus and E. atrocostata, the largest localized population
declines of other species are associated with the developed (resort) section of the atoll. This
may be related horticultural / architectural practices that restructure typhoon adapted
vegetation allowing complete overwash and loss of most structures, soil, and sand during
severe storms.
Considering the absence of all lizards on the resort 1/3 of the atoll, a substantial fraction
of the lizard population was lost because of the cumulative effects of typhoons. Habitat
5
specialists E. atrocostata and C. poecilopleurus probably suffered much greater population
declines, which is related to susceptibility of these habitats to typhoon damage. Despite
that, the data suggest that relatively undisturbed atolls will tend to retain herpetofaunal
components despite substantial typhoon influences.
ACKNOWLEDGMENTS
Assistance in the field was provided by Rebecca Hensley, Robert Cruz, and Earl
Campbell III. Gordon Rodda and Thomas Fritts generously provided unpublished data.
Rebecca Hensley, Gordon Rodda, Thomas Fritts, and Kevin de Queiroz commented on a
version of the manuscript. Portions of this study were funded by the Endangered Species
Conservation Program, Project E-4 (to Guam) and by the U. S. Department of the Interior,
National Biological Survey.
LITERATURE CITED
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Guam, Pacific. NOAA Natl. Clim. Data Center, Asheville, NC. 8 p.
HENSLEY, R. A. and M. J. MCCOID. 1994. Cryptoblepharus poecilopleurus (Snake-
eyed Skink). Activity. Herpetol. Rev. 25:121.
JACKSON, W. B. 1967. Productivity in high and low islands, with special emphasis
to rodent populations. Micronesica 3:5-15.
MCCOID, M. J. 1993. The 'new' herpetofauna of Guam, Mariana Islands. Herpetol.
Rev. 24: 16-17.
.and R. A. HENSLEY. 1994. Distribution and abundance of Perochirus ateles
(Gekkonidae) in the Mariana Islands. Herpetol. Rev. 25: 97-98.
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slevini (Scincidae) in the Mariana Islands. Herpetol. Rev. 26: in press.
MYERS, R. F. 1991. Micronesian Reef Fishes. 2nd ed. Coral Graphics, Barrigada,
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NEUBAUER, C. P. AND D. R. NEUBAUER. 1981. The vegetation of Cocos Island
(Mariana Islands). In L. Raulerson (ed.). Plant biogeography of Guam. Univ.
Guam Mar. Lab. Tech. Rep. 69. pp. 23-39.
RODDA, G. H, T. H. FRITTS, AND J. D. REICHEL. 1991. The distributional
patterns of reptiles and amphibians on the Mariana Islands. Micronesica 24: 195-
210.
, M. J. MCCOID, AND T. H. FRITTS. 1993. Adhesive trapping II. Herpetol.
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2p.
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ATOLL RESEARCH BULLETIN
NO. 440
FLOWERING AND FRUITING IN THE FLORA OF HERON ISLAND,
GREAT BARRIER REEF, AUSTRALIA
BY
R.W. ROGERS
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
APRIL 1996
yi \ Genie? See
WITAIIUG HORAGEAS JI01 ay
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ove) JADA
FLOWERING AND FRUITING IN THE FLORA OF HERON ISLAND,
GREAT BARRIER REEF, AUSTRALIA.
BY
R.W.ROGERS'
ABSTRACT
The plant species in flower and fruit on Heron Island, a sandy cay on the Great Barrier
Reef, Australia, were observed at intervals of three months for three and a half years. At no
time were less than 20 nor more than 36 of the 49 species monitored found to be in flower, nor
were less than 20 nor more than 41 of the 50 species monitored found to bear fruit. Despite a
strongly seasonal climate there was not a strong seasonal pattern evident in the number of
species in flower or fruit, although some species were themselves strongly seasonal. A principal
components analysis of all flowering records, however, demonstrated a seasonal polarity with
March and September representing the two extremes. Fleshy fruited species, important for
frugiverous birds such as silvereyes, bore fruit throughout the year.
INTRODUCTION
Temporal patterns in flowering and fruiting are significant attributes of vegetation, for these are
attributes subject to selection as are any others. Variation in seasonal flowering patterns has
proved to be significant in understanding of heathlands in Australia (Specht et a/. 1981) and
Europe (Woolhouse & Kwolek 1981), both in terms of ecophysiology, and in terms of the
evolutionary derivation of the floras. The availability of flowers and fruit is manifestly
important to those animals which depend on fruit, seed and nectar as food resources, and an
interaction between plant phenology and the birds responsible for seed dispersal has been
postulated (Herrera 1986).
There has been little previous study of the temporal variations in flowering and fruiting
of the plants on the cays of the Great Barrier Reef, although Heatwole (1981) noted flowering
times for a few species on One Tree Island. There has apparently been little if any study of
seasonality on similar islands elsewhere. Opportunity to visit Heron Island on a regular basis
was therefore used to collect information on the flowering and fruiting patterns of the flora of
that island.
Heron Island is a small coral cay about 800 m x 300 m situated 70 km offshore from
the Australian continent (Lat. 23°26' Long. 151°5S'E). Much of the island is clad in a dense
forest of Pisonia grandis trees, with some fringing grasslands (Fosberg 1961). The island has
been the subject of extensive weed invasions during its recent
history of intense human activity, and this has been documented in particular for a spoil dump
established on the island in 1987 (Rogers 1993) and in more general terms by Chaloupka &
Domm (1986).
‘Botany Department, The University of Queensland, Queensland 4072, Australia.
Manuscript received 4 May 1994; revised 21 April 1995
Heron Island has a strongly seasonal climate, with mean rainfall varying from as little
as 20 mm in September to as much as 145 mm in February. The four months July to November
receive in total less than 15% of the annual rainfall (Fig. 1). A consequence of this strong
seasonality is that Pisonia grandis and Ficus opposita often lose many of their leaves by
December, and the native grass cover of the island dies.
160
140
120
100
80
60
Mean rainfall (mm)
40
20
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Month
Figure 1 Distribution of mean monthly rainfall through the year for Heron Island (from Walker
1991a).
Methods
Heron Island was visited at intervals of about three months between September 1990 and
December 1993, and the flora surveyed to determine which species were represented by
individuals in flower or fruit at the time. These data were recorded and analysed in terms of
total species in flower and fruit, native plants in flower and fruit, weedy plants in flower and
fruit, native dicots in flower and native and weedy grasses in flower, and numbers of species
bearing fleshy fruit.
Information was collected in September, December, March and July, for convenience
3
referred to as spring, summer, autumn and winter. Records were made by walking a series of
transects on the island which traversed all the vegetation types and the habitats of all the plant
species known for the island.
To detect pattern in a multivariate system such as that studied here where it is proposed
to detect seasonality of flowering in a system comprising 49 species requires a multivariate
analysis. One such analysis which is suitable for the purpose is Principal Components Analysis,
and in order to reveal underlying patterns in the data a principal components analysis was
executed on the flowering data (Wilkinson 1990).
Table 1 Seasonality of flowering of plants on Heron Island, Great Barrier Reef, Australia,
between September 1990 and December 1993. N = total number of plants in the category.
Number of species in flower
year 90 90 91 91 91 91 92 92 92 92 93 93 93 93
Month Si) De Mads) Die Med Sab) YM.) Si iD
N
All species
natives 2X5 ALO) 4s IS ass ah aL fs} La US) LG L114) aly 9)
weeds AS} 6 iO) WS 7 US) a7 12 WS) Qi 16 14 1 2B 22
Total 49 28 24 32 35 30 34 20 30 36 32 26 33 40 41
Dicots
Native 7x 0) 13} Ws 1G 12 16 7 YF Ws les ail 12 WH il6
Weeds TS 7) 9S Be pelea 7S a nO MO, 2 aS) aS
Total 2) 22 20) 23 2y 2A Bil 14 22 2 2 ZO AX BO Sil
Grasses
Native Vat dL, wR TUITE 2), Geen ORES My goal aOR (2) dialer er Lee On eA ee
Weeds Sh See Sy ime GwG. GS.) Cho ei wig eae 7M iBict 7
Total ib 6 4 SF Bo 6 FW. 6 tS) th By) 8) ao). ako)
Table 2 Seasonality of fruiting of plants on Heron Island, Great Barrier Reef, Australia,
between September 1990 and December 1993. N = total number of plants in the category.
Number of species in fruit
year SOR90 NST eS 1 Oi 91929269259 21493593.9393
Month Sap Oto. DOM ed So. DOM wd. 6S. 6D
N
All species
natives PAI) Wyo De Th ALS) PAO) AUG) AUS) TLD oll Ah ENT ood E'S) TL fg ILS} hI IAS Dae
weeds 28) Gh MoD may iG Oe ISteGw20Na Tei A TS 220
total 50,29..26536936136 .40,,20,.30)\37,36. 29..37:1:40 41
Fleshy fruits
Att AAT EREUGARAMA AMAR YAN 41S edie Bit &
45
40
Species in flower
S90 $91 $92 $93
Season
Figure 2.The total number of plant species in flower (4), number of native species in flower
(CQ), and number of introduced species in flower(™) Heron Island, Great Barrier Reef, Australia,
at three month intervals from September 1990 until December 1993.
RESULTS
Flowering was recorded for 46 of the 49 species examined for flowers (appendix 1):
Ipomoea pes-caprae and Salsola kali were not seen in flower, and flowering of Ficus opposita
was not determined because of the syconium within which flowers are produced.
No distinct patterns of number of species in flower around the year was apparent (Fig. 2, Table
1), whether the flora was taken as a whole or divided into components. The patterns of
variation shown by all groups of species are essentially the same. A depression in the number
of species in flower is apparent in July (southern winter) of 1992 and 1993, but July 1991
shows a high number of species flowering.
Patterns can be detected in individual species (appendix 1). Apiwm leptophyllum,
Bromus catharticus and Wollastonia biflora flower only in spring-summer, whereas Cordia
subcordata may be found flowering in any season except summer. Pisonia grandis flowers in
summer-autumn. The small herb Gnaphalium luteoalbum appears to flower in any season but
autumn. Of the remaining species 17 were recorded flowering in 12-14 of the times surveyed,
and 4 in three or less occasions.
ar iN eet aE iin a
5
The principal components analysis showed that over 50% of the variance in the
flowering matrix was explained by the first two components (36.3% and 15.4% respectively),
and that the replicate seasonal collections for March and September represented poles of an
ordination, with the summer and winter collections (December and June) falling into an
intermediate position (fig.2)
Component 2
6
0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9
Component 1
Figure 3. Plot of the first two components from a Principal Components Analysis of three-
monthly flowering records from Heron Island. Each record is identified by month (M = March,
J = July, S = September, D = December) and year.
Fruiting was observed in 47 of the 49 species (appendix 2), Ipomoea pes-caprae and
Salsola kali not being observed in fruit during the study period: expanded syconia of Ficus
opposita were treated as fruits. No distinct seasonal patterns were evident in numbers of species
fruiting around the year, and the total number of species in fruit closely paralleled the number
flowering (fig.3). At least four fleshy fruited species were in fruit at any time, and three of
these, Tournefortia argentea, Ficus opposita and Pipturus argenteus bore fruit every time they
were observed.
DISCUSSION
Observations made four times per year do not permit a study of seasonality of flowering
in any great detail. However, it is sufficiently frequent to detect the more striking patterns
6
which may be present. It is apparent that the very strong seasonality evident in the rainfall
pattern for the Island is not reflected in the number of species in flower or fruit. The polarity
demonstrated by the principal components analysis between March and September, however,
is quite clear. Thus, although the seasonal differences are diffuse, involving numbers of species
showing quite different patterns, the overall view is of well developed seasonality in flowering
pattern.
At all times studied there was a higher proportion of weed species in flower than of
native species, and at all times except December 1993 a higher proportion of weed species in
fruit than native species. This is similar to the finding of Odgers and Rogers (1993) that weedy
grass species growing together with native Australian grass species showed longer flowering
seasons than did the native species. It is not surprising that weedy species, characteristic of
frequently disturbed sites, are more likely to have individuals in a reproductive condition than
are those species which characteristically occupy relatively undisturbed natural systems.
Some of the native species, however, flower and fruit continuously. The continual
fruiting of Ficus opposita, Pipturus argenteus and Tournefortia argentea, all of which have
fleshy structures associated with their fruits, is probably of great importance to the resident
population of silvereyes (Zosterops lateralis) and bar-shouldered doves (Geopelia humeralis)
on the island, indeed, Walker (1991b) ascribed the extinction of bar-shouldered doves on Lady
Elliot Island to the loss of Ficus opposita from the flora of that island. It is also likely to be
important for the plants that a population of seed dispersers is maintained. A continuous
flowering is probably also important to the plants in that it permits the maintenance of
populations of pollinators in an otherwise isolated habitat. The nectar associated with flowering
may also be important to the silvereyes (Barker & Vestjens 1989) and raises the possibility of
bird pollination in Tournefortia argentea which has strongly nectiferous flowers, and perhaps
in Pipturus argenteus too.
The strongly seasonal native trees Cordia subcordata and Pisonia grandis appear not
to depend on permanently resident birds for seed dispersal. P. grandis seeds are commonly
associated with the migratory sea bird populations which nest amongst them during the fruiting
season, especially the white-capped noddy (Anous minutus), and are presumably dispersed
adhering to birds (Walker 1991a), although the noddies may suffer heavy fatalities from high
P. grandis fruit loads in those years in which P. grandis fruiting is heavy. Cordia subcordata
produces a large and rather corky fruit which is often found in drift along beaches and is
apparently dispersed by flotation.
Boerhavia tetrandra is in the same family (Nyctaginaceae) as Pisonia grandis, but is
a prostrate herb which bears sticky fruit similar to that of P. grandis, but displayed only a few
centimetres above ground level. Seedlings of Boerhavia tetrandra are commonly observed in
disturbed areas of Heron Island, whereas those of Pisonia grandis are seen very rarely, but then
in very large numbers within the forest (e.g. March 1972 when several hundred could be found
in a square meter). The sticky fruit of Boerhavia tetrandra, however, are more likely to be
dispersed by ground feeding birds such as the doves and rails, in contrast to the dispersal of
Pisonia grandis seeds by marine bird species.
The possibility of coevolution of the frugivorous birds and the flora cannot be
dismissed, especially in terms of selection for plants which have an extended fruiting season,
for Herrera (1986) has observed that changes in phenology are amongst those most likely to
occur in response to frugivory and contingent dispersal. It is recognised that the silvereyes of
the Capricorn Group are a distinct variety, differing from their mainland relatives. However, it
is not possible to argue for any close co-evolution between the birds of the cays and the flora
in the way in which Reid (1991) argued for coevolution of mistletoes and mistletoe birds,
although in arid zones mistletoe birds may be dependent on very few species of mistletoes for
7
survival, just as frugivores are on a coral cay. Comparative studies of the eating habits of the
mainland and cay varieties of silvereyes and of the seasonality of flowering in Tournefortia
argentea may be profitable in this context.
Acknowledgements
I am grateful to the Queensland National Parks and Wildlife Service for permission to work on Heron
Island, and to the staff of the Heron Island Research Station of the University of Queensland for
assistance.
REFERENCES
Barker, R.J. & Vestjens, W.J.M. 1989. 'The Food of Australian Birds. 2. Passerines'. CSIRO,
Canberra.
Chaloupka,MY., and Domm, S.B. 1986. Role of anthropochory in the invasion of coral cays
by alien flora Ecology 67, 1536-1547.
Fosberg, F.R. 1961. Description of Heron Island. Attol Research Bulletin 82, 1-4.
Herrera, C.M. 1986. Vertebrate dispersed plants: why they don't behave the way they should.
In 'Frugivores and Seed Dispersal' (eds A Estrada & T.H.Fleming) pp 5-18. Junk,
Dordrecht.
Odgers, B.M. and Rogers, R.W. 1993. Contrasting diaspore and vegetation attributes from
natural and disturbed habitats in an urban eucalypt forest reserve. Australian Journal
of Botany 41, 637-648.
Reid, N. 1991. Coevolution of mistletoes and frugiverous birds. Journal of Ecology 16, 457-
69.
Rogers, R.W. 1993. Plant colonization of a rubble bank on Heron Island, Great Barrier Reef,
Australia. Atoll Research Bulletin 384, 1-8.
Specht, R.L., Rogers, R.W. & Hopkins, A.J.M. 1981. Seasonal growth and flowering rhythms:
Australian Heathlands. In 'Ecosystems of the World 9B: Heathlands and Related
Shrublands' (ed R.L. Specht) pp 5-13. Elsevier, Amsterdam.
Walker, T.A. 1991a. Pisonia Islands of the Great Barrier Reef. Part 1. The distribution,
abundance and dispersal by seabirds of Pisonia grandis. Atoll Research Bulletin 350,
31-39.
Walker, T.A. 1991b. Pisonia Islands of the Great Barrier Reef. Part 3. Changes in the vascular
flora of Lady Musgrave Island. Afoll Research Bulletin 350, 31-39.
Woolhouse, H.W. & Kwolek, A.V.A. 1981. Seasonal growth and flowering rhythms in
European heathlands. In ‘Ecosystems of the World 9B: Heathlands and Related
Shrublands' (ed R.L.Specht) pp 29-38. Elsevier, Amsterdam.
Wilkinson, L. 1990. SYSTAT: "The System for Statistics'. Evanston, Systat Inc.
8
Appendix 1: Flowering calendar for Heron Island.
1 = plant seen in flower; 0 = no plant seen in flower.
Year 90) 9091 191 91194" 92192 92%92)\93).98. 93493
Month S cD. pM AS Dow. oT Si) Di iS ED
Species
Abutilon indicum
Achyranthes aspera
Amaranthus viridis
Apium leptophyllum
Bidens pilosa
Boerhavia tetrandra
Brachiaria
subquadripara
Bromus catharticus
Cakile edentula
Calyptocarpus vialis
Capsella
bursa-pastoralis
Cassytha filiformis
Casuarina
isetifolia
Celtis paniculata
Cenchrus echinatus
Commicarpus insularum
Conyza bonariensis
Cordia subcordata
Coronopus didymus
Cynodon dactylon
Digitaria ciliaris
Eleusine indica
Euphorbia atoto
Euphorbia cyathophora
Euphorbia prostrata
Euphorbia tannensis
Ficus opposita
Gnaphalium
luteo-album
Ipomoea pes-caprae
Lepidium virginicum
Malvastrum
coromandelianum
Pandanus tectorius
Pipturus argenteus
Pisonia grandis
Poa annua
Portulaca oleracea
Salsola kali
Scaevola taccada
Sisymbrium orientale
Solanum americanum
Sonchus oleraceus
Sporobolus virginicus
Stenotaphrum
micranthum
Suriana maritima
Thuarea involuta
Tournefortia argentea
Tribulus cistoides
Wollastonia biflora
(je)
SOPPORHP OPRPERPOOROPROO POP BROOPRHPORHPOOHOOO OF OFPHO BHRHPHPO
PRERPRPRP OCOFPOPOOORPOFP HOO BPROORPHFPOFPACDD0D00GO CF COCO HORHOP
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COPPRPERRP OFPFPRPOCORPORPORP POP BEBBBPBEBBPEBHOHROHOO OF BHOPR BROPREPE
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PREPRPRPRP PRBPRPRPORPORPRPORP POR BRBBBBBPBBBHBHBEHBHO CO BHBBHP BPRPRBPHPOP
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a. =
Appendix 2: Fruiting calendar for Heron Island.
1 = plant seen bearing fruit; 0 = no plant seen bearing fruit.
Year 90 90 91 91 91 91 92 92 92 92 93 93 93 93
Month Se'DO MiG Ss) DM U.S DM UJ S D
Species
Abutilon indicum i
Achyranthes aspera
Amaranthus viridis
Apium leptophyllum
Bidens pilosa
Boerhavia tetrandra
Brachiaria
subquadripara
Bromus catharticus
Cakile edentula
Calyptocarpus vialis
Capsella
bursa-pastoralis
Cassytha filiformis
Casuarina
equisetifolia
Celtis paniculata
Cenchrus echinatus
Commicarpus insularum
Conyza bonariensis
Cordia subcordata
Coronopus didymus
Cynodon dactylon
Digitaria ciliaris
Eleusine indica
Euphorbia atoto
Euphorbia cyathophora
Euphorbia prostrata
Euphorbia tannensis
Ficus opposita
Gnaphalium
luteo-album
Ipomoea pes-caprae
Lepidium virginicum
Malvastrum
coromandelianum
Pandanus tectorius
Pipturus argenteus
Pisonia grandis
Poa annua
Portulaca oleracea
Salsola kali
Scaevola taccada
Sisymbrium orientale
Solanum americanum
Sonchus oleraceus
Sporobolus virginicus
Stenotaphrum
micranthum
Suriana maritima
Thuarea involuta
Tournefortia argentea
Tribulus cistoides
Wollastonia biflora
kh
OPFPOOO OCOPREFPRPODOORPOPRRO POP BPREROORPORHOOHHOH OF OFPBPH BPRELHO
PRROPF OCOFPPOOOOORPBRHE POP BEOORPPORPOCOHODOH OH OHOO HOOrOH
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PRROPP OPRPEPRPORPRPORBRHEP COP BPROPRPOPOPBPBPBPBBEHP OF OFFPO BREEHPOPE
PRRPOPP OPRPREPRPORPRPRPRPEBEP POP BRRPRPOPBPEBHPORPOPRHBPHE BPH OPRHPO BPREBPHPOPE
SCPFPFPODOD BPODDDODDCOOPRHBHP COO BEBPRPORRPEOOFPOPEHEHR CO C000 HBOOROLH
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ATOLL RESEARCH BULLETIN
NO. 441
NAMU ATOLL REVISITED: A FOLLOW-UP STUDY OF 25 YEARS OF
RESOURCE USE
BY
NANCY J. POLLOCK
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
APRIL 1996
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The Marshall Islands
NAMU ATOLL REVISITED:
A FOLLOW-UP STUDY OF 25 YEARS OF RESOURCE USE
BY
NANCY J. POLLOCK
ABSTRACT
Reliance on local resources for food remains even though the population of Namu has
doubled in size. Breadfruit, pandanus and fish are still the main subsistence foods on an atoll in
the northern Marshalls. A restudy of household food uses 25 years after the original study
revealed few changes in the supply, but dependency on those foods had doubled, thereby
increasing the risks from both natural and market failures. Marshall Islands government support
will be necessary to maintain such a population on their home atoll.
INTRODUCTION
Atolls are ever-changing environments. Nature changes them with all her complex
interactions. Human activity also brings many additional changes. This paper documents the
impact of human lifestyle over 25 years on one islet of an atoll in the Northern Marshal Islands.
It aims to add to the work of a number of scientists who have studied the Northern Marshal
Islands over a forty year period. The paper is a vote of thanks to Dr. Ray Fosberg whose work
contributed greatly to my attempts to understand the interaction between people and their island
environment, particularly their food resources.
Data on food resources were gathered over a period of fifteen months spent on Namu Atoll
in 1968 and 1969 with the aim of understanding the interrelationships between food use and
social organization (Pollock 1970; 1992). Similar data have been the subject of a year long study
during 1992-3 in the Marshal Islands, that enabled me to pay two return trips to Namu for a
week each, one in November 1992 and another in August 1993. Most of my work has
concentrated on one islet, Majkin on the eastern side of the atoll.
Namu lies in Ralik or the western chain of the Marshal Islands, some 25 miles southwest of
the large atoll of Kwajalein. The lights of Kwajalein can be seen from the northern shores of
Namu Namu islet, the northernmost piece of land in Namu Atoll. There are some 56 islets in
Namu Atoll, only three of which are regularly inhabited. Namu Namu was the first settled,
according to local legend then Majkin to the eastern side, and finally Mae-Leuen in the south.
The latter is two islets joined together by a sand bar that formed after a Japanese ship wrecked
on the reef during World War II. The whole of Namu Atoll stretches over some 25 miles from
north to south, and is 15 miles at its widest point.
Dept. of Anthropology, Victoria University, Wellington, New Zealand
Manuscript received 25 February 1994; revised 13 December 1994
Rainfall amounts to some 80" - 120" p.a., so that Namu lies near the northern limit of
adequate moisture for breadfruit trees to flourish. Today they dominate the upper canopy, as seen
from the air, with a few old Lukwej (Calophyllum inophyllum) trees reaching similar heights.
The coconut trees are tall too, but are becoming spindly, as many are now 50 and 60 years old.
The atoll has been hit by several cyclones in recent years, the latest in December 1992 caused
considerable damage to the vegetation; FEMA aid was provided to the people of Namu.
Each islet is divided by the Marshallese into three use zones, lik or oceanside, eolap or
middle of the island, and iar, or lagoonside. These three zones refer to particular kinds of
vegetational zones as determined by human activity. These activities have resulted in differential
soil fertility. The usage of these zones is determined in part by the orientation of the piece of
land to wind and waves. In addition each islet is divided into two parts, jittoen and jittoken,
terms that originally referred to the ways a canoe could head.
The lik side of Majkin is exposed to the prevailing northeast wind and thus is battered by salt
spray, even though there is a wide protecting reef. Hence only salt tolerant strand vegetation such
as Scaevola or Morinda citrifolia (nin) grow here. This side of the islet is mainly used by adults
for defecation on the reef; that usage places taboos on visiting the area for other reasons. Eolap,
or the middle of the island is quite a small area at either extremity of Majkin, but in the centre
of the atoll this area can cover a quarter of a mile. Here the greatest proportion of the coconut
trees have been planted, and pits excavated to grow the form of taro best suited to atolls, namely
iarej, Cyrtosperma chamissonis. Pandanus trees in a number of varieties, have also been planted
on the lagoon side of this area mainly for use of the leaves in making handicrafts.
Lar, or the lagoon side of the island, is the most heavily used part of the atoll. Residences,
the road and useful trees are concentrated here. The main reason is that this area gives best
access to the fresh water lens accessed by means of wells. This fresh water supply is also tapped
by the breadfruit trees which have been planted around the residence areas both to give shade
and to provide fruit and leaves. Residential areas are readily distinguishable by the white coral
which is gathered from the lagoon shore to spread around the house sites to keep them clean and
improve drainage. The lagoon shore is also the landing site for any visiting ships, whether sailing
canoes, one man dug-out canoes, faster launches with outboard engines, or occasional deep
water field trip ships.
Landholdings reflect the tripartite use of the land. Each landholding or weto, extends from
oceanside to lagoonside in more or less straight lines. This means that the landholders can make
best use of the various parts of the land. Each weto is named and is controlled by a particular
matrilineage. Members of that matrilineage have the rights to live there and to make copra on
that land and to plant new vegetation. There are fourteen named weto on Majkin, each running
across the island; they vary in width, some being very wide. The two weto at either end of the
islet are not as productive as those in the middle of the islet because of limited access to fresh
water.
The core social group that shares a residence weto on Namu is two or more sisters and their
descendants, together with their nuclear families. That group forms one household. They may
have several sleeping houses, but they share a common cookhouse where two or three women
3
of the group cook for everyone. Households on Majkin vary in size from 12 to 75 people, even
getting as large as 110 when special events, such as a church conference, are held on this islet.
Chief decision maker for a weto , known throughout the Marshal Islands as the alab, is
usually the oldest brother of the lineage. He may not be resident in the particular household he
controls but is usually resident on the islet at another weto. He manages all affairs connected
with the plot of land, including planting new crops and digging wells and erecting new
structures. He thus is an important influence on the productivity of a weto. In 1968 when the
whole islet was short of food and one weto group had none at all, a comment was made that their
alab in times past had not planted well for their lineage, and thus he was to blame for their plight.
The main starch food sources on Namu are breadfruit and pandanus, with some arrowroot.
These are eaten with some fish or shellfish or coconut as an accompaniment (jalele). Rice is
heavily used, being purchased from field-trip ships with money earned from the sale of copra.
Some flour is also purchased to make a type of loaf, or dumplings. The pattern of food use has
not changed drastically over the twenty five year period as discussed below, even though the
population of the atoll has doubled. It has just become more dependent on rice to balance the
supply when there is little or no breadfruit or pandanus.
Breadfruit
Breadfruit trees dominate the house sites. These are mainly the seedless variety (Artocarpus
altilis) of the Bitaakdak, Bukdol/Bukarel varieties, though seeded varieties (Artocarpus
mariennensis) such as Mejwaan are also planted. A house by house assessment of breadfruit
trees on Majkin in July 1993 revealed an average of 6 trees per house, with one house having
27 trees and another only 4. Most of the breadfruit trees were 60 to 70 feet tall and had a base
trunk circumference of between 8 and 12 feet. They grow around the residence site with at least
one being fairly central to provide shade. A few smaller trees have been planted more recently
by taking a cutting from the root of an old tree. These young plants are carefully nurtured until
they can stand alone.
People on Majkin cannot remember when particular trees were planted, but judging by the
similarity in size of the largest trees, there must have been a concerted effort to propagate new
trees in the 1930s to 1950s, but after that few new trees were planted until the late 1980s. As result
food resources will be severely curtailed when those older trees pass the peak of their bearing
lifespan. In 1993 most of them were bearing heavily.
Breadfruit trees are so valuable as a food source for the whole residence group on one weto
that one is cut down only out of extreme necessity. Indeed the Marshallese term for breadfruit,
ma, is also the generic Marshallese name for a plant, indicating that breadfruit has prime status
in their categorization of plants. In former times a tree might be cut down if a canoe was sought
by the paramount chief for a particular reason.
The green, globular fruit of the seedless variety is round like a boy's head, as Dampier (1616)
described it, and weighs on average 3 to 5 Ibs. On Namu, a breadfruit tree will produce fruit
three times a year in a good season, that is, one without drought or cyclones. The main season
is May through August when the greatest number and largest fruit are produced. Two secondary
4
seasons of fewer and smaller fruit do occur in October/November and January/February. The ten
or so different varieties mature at slightly different times, thereby lengthening the breadfruit
season.
The fruit of the seeded variety, Mejwaan, is very different from the seedless variety. It is of
irregular shape and weighs about 2 or 3 pounds. One crop matures in late April and May, and
may have a small second season. Both the pulp and the seeds are eaten. The flesh has a slightly
more tangy flavour than that of the seedless breadfruit (NJP Namu fieldnotes 1968) (For a full
discussion of breadfruit varieties throughout the Pacific, see Ragone 1987).
Namu people cook the seedless variety in a number of ways. The most common way is to
roast the whole fruit in the coals and then scrape off all the charcoal before serving (kwonjen).
Breadfruit may also be boiled and coconut cream added at the end to make a dish called
bwilitudek, or baked in the earth oven as a whole fruit to which coconut cream has been added
in the centre(beljij). Another five ways of cooking breadfruit were recorded during the 1993
season.
At the end of the season, particularly the main season, the ripe fruit are picked just before
they are ready to fall in order to make them into fermented breadfruit paste (bwiro). The process
involves all the families associated with a household, even those living elsewhere but strongly
attached such as a brother or sister; even 8 or 9 year olds and those in their 70s help in the
peeling stage. The fruit is first peeled, then quartered, and placed in sacks to be soaked in the
lagoon for several hours. Then it is left for two nights resting in a tree to drain and begin the
fermentation process. Once fermentation has commenced and the fruit has become mushy, four
or five sacksful are tipped into one pit. These pits are made by excavating a hollow in the sand
and lining it with old breadfruit leaves; each pit is then covered with old breadfruit leaves and
weighted down with several coral slabs. These storage pits are usually placed near the
cookhouse. The paste stays there fermenting for a month or more before it is ready to be
processed for eating.
Each household on Majkin was very active making bwiro in early August 1993. Some
families had filled five or six pits already and expected to add still more. The people were
rejoicing because the season had been so good. By October and November they will begin to use
the fermented paste for their daily food supply, and also send some of the loaves baked from the
paste away to their relatives in Kwajalein.
On other atolls in the Marshal Islands, however, breadfruit is being allowed to fall and rot
and is not being made into bwiro. Laura, at the western end of Majuro Atoll, is one such islet that
has a multitude of breadfruit trees and yet the fruit are being wasted. There are two possible
reasons; firstly the people prefer store food because it is quicker to prepare, and secondly money
is more readily available as many households have one or two persons working for wages at the
other end of the atoll. The Ministry of Resources and Development has therefore accepted
external aid money to build a factory in Laura to make breadfruit chips. These will be made from
the fruit at the height of the season, those people owning several trees selling the fruit to the
factory. The chips will be marketed like potato chips in small packets as a snack food. Such
projects have been successful in Western Samoa and other parts of the Pacific. The project will
5
thus utilize a resource that is being wasted at the moment, and also produce something locally
as a Substitute for an imported product. The managers of the project aim to extend utilization
of the plant for the same chipping process with other crops such as iarej and bananas; these snack
foods will be sold locally and exported.
Besides its fruit, the breadfruit tree has several other uses. Its leaves, both green and brown
are used extensively for wrapping food to be placed in the earth oven, and to cover the earth
oven before earth is piled on top. The leaves can be used as an instant plate, and to cover food
left in a container. The sap of the breadfruit tree is a well known form of glue used in calking
canoes and in handicrafts. The trunk was formerly the most favoured wood from which to make
a dugout paddling canoe (korkor) or to make planks for the larger sailing canoe (tibnil). Few
trees have been cut down recently, however, fishermen preferring to cannibalize old canoes to
patch up one that is broken. Dead wood from breadfruit and other trees contributes to the fuel
supply. The detritus including leaves forms valuable mulch on land that has very little humus.
Pandanus
The pandanus tree (bob) is much smaller than the breadfruit, standing only some ten or
fifteen feet high at the most. Its distinctive feature is its prop roots which in an old tree may be
three or four feet long. Two distinct varietal groups have been propagated over time to meet local
needs on an atoll such as Namu, one for the production of fruit, and one that produces the best
leaves for making handicrafts. The two uses mean that new varieties are selected on the basis of
their appropriate qualities. Many varieties are named by local people, and their attributes clearly
distinguished. In 1968 I recorded 24 different varieties growing on Majkin; in 1993 | was told
that double that number now exist, though | did not record the names.
Pandanus fruit are large and globular. They weigh some 20 to 35 lbs. Each fruit consists of
some 50 or 60 drupes that are attached to a central stem. Each drupe has a hard generally green
exterior and a fibrous ‘brush’ interior surrounded by an orange pulp when ripe. The fruit bearing
qualities of the pandanus have been carefully selected for by atoll populations across the Pacific;
for other Pacific societies the pandanus is considered rubbish food. The plant is grown by
vegetative propagation using a slip from one of the prop roots. The fruits seldom contain seeds,
and if they do, the resulting plant will not be fruit-bearing.
The pandanus season on Namu begins in August or September and lasts through to
December. The fruits are eaten when ripe by breaking off a drupe from the central stem and
rubbing the fibres between the teeth. Other varieties may be cooked to soften the fibres. The
process of eating pandanus resembles sucking on a shaving brush. Since the edible part is so
fibrous, the eater ends up with many strings between the teeth. The paste is high in vitamin A
and thus is a valuable addition to the diet.
Pandanus is not considered a main source of food by the Marshallese, though it is eaten
extensively in season. Rather it is used as a snack by both adults and children alike. Formerly
the pulp was extracted from the fibres by pressing them against a v-shaped object made of shell,
wood or (today) metal. This juice was then boiled until it thickened, and the paste set out on
mats to dry in the sun for three or four days. This dried product was rolled and tied to form a
6
product known as mokwan; it was carried by sailors as it did not deteriorate in its leathery form.
Such preservation is seldom carried out today as it is time consuming and there are more
convenient imported foods.
An alternative recipe was to add arrowroot starch to the boiled pandanus paste, plus some
coconut cream to yield a food known as peru. This was considered a delicacy and so was only
made on special occasions, or as a gift for the paramount chief. It required a lot of time to
prepare. It did not keep, so was not a form of preservation. Today it is made only very
occasionally.
A second major use of the pandanus tree is in the manufacture of handicrafts (amimono).
The green leaves are cut, dried and processed into strips which can be woven into sleeping mats,
or smaller objects, or are boiled to produce a very fine white fibre for special basketry.
Alternatively the dried leaves, once the spine is removed, may be processed into coarser mats,
or into thatch.
Handicrafts have become such a mainstay of the economy that considerable effort has gone
into finding the right plant for the particular product desired. This trade has led to a
diversification in the species of pandanus grown.
Fishing
Fish are the third major resource which is heavily utilized on Namu. Fish are considered a
highly desirable complement to the starch portion of the daily meal, but on Majkin they are are
a luxury. They are not easily caught, and with a large population to be fed there are never enough
to satisfy everyone. In part this shortage is due to difficult access across the reef on the oceanside
of the islet, and in part to the shortage of fish on the lagoon side. Whether the latter is due to
over-fishing, or some ecological anomaly is not clear.
Every household aims to have some fish for the Saturday evening meal. So those men who
have access to some form of boat, whether one-man dug-out or a launch with an outboard motor,
spend the day fishing in the hopes of providing a decent supply for everyone in the household
to have a small portion on Saturday night, and hopefully some left over for Sunday. But they are
not always successful, so that fish is available for only about half the weekends of the year
(Pollock fieldnotes 1968.).
The greatest amounts are caught by those who use the launch to go farther out into the
lagoon, or even out the pass on the other side. But that requires gas, and gas is a scarce
commodity. It can only be purchased from fieldtrip ships, and then only five or six drums at a
time. So frequent trips across the lagoon are not possible and men tend to fish close to the lagoon
shore. Even the good fishermen will tell you they can sit there for hours and catch only four or
five fish. But when the launch goes out they will catch some 30 or 40 Ibs of fish. The people of
Majkin would like to have a more regular supply of fish.
A new fisheries facility (built by MIMRA) has just been completed in the centre of Majkin
islet right on the lagoon shore. It consists of storage facilities, a fresh water tank, two large
launches, and a tractor for launching them. This is one of three such facilities already built in the
Marshalls with aid money to assist the people to become more self sufficient in fish. The aim
i
is to provide the facilities for catching fish for their own needs, and also to catch fish to sell to
Ebeye, the urban concentration on Kwajalein. In August 1993 the project was awaiting the
appointment of a director to begin operations.
It will be interesting to see how successful this operation is, given the ongoing difficulties
of catching fish on Majkin. In contrast, Namu Namu islet to the north of the atoll has a plentiful
supply of fish as they can use both the ocean side as well as the lagoon side of their islet. On the
ocean-side they catch flying fish (jojo) in season. Thus it is surprising that MIMRA (Marshall
Islands Marine Resources Association) did not see fit to build the new fisheries project plant at
Namu Namu. Time will tell.
Continuities and Changes
These basic resources have remained unchanged. What has changed is the demands upon
these resources. The population of Majkin islet has doubled to about 440 people from 200 in
1968. People move constantly between the three islets of the atoll, and also out beyond the atoll,
so it is difficult to give an exact figure at any one time. The household survey we conducted in
late October 1992 yielded a total of 517 people, but a similar survey in August 1993 yielded a
total of only 347. The reason for the large number in 1992 was that many people had travelled
from Namu Namu and Mae-Leuen, and some from Ebeye to take part in a church conference on
Majkin. In addition, the numbers in August 1993 were down by about 50 people who were away
at another church conference in Majuro during July 1993. So the demands of the population on
the resources do fluctuate quite considerably. Tree crops and fish are admirably suited to such
irregular demands.
Copra is still the main source of cash as it was in 1968. The price of copra as paid to the
producer has fluctuated considerably over the years; it was so abysmally low in 1968 at 2 cents
per pound that Namu people (along with other Marshallese) were seriously questioning
whether it was worth the effort. But without copra money they could not buy rice, flour, tea and
sugar, so it was better than nothing. In 1990 the Marshallese government agreed to support the
price of copra at 15 cents per pound in an effort to draw people away from the urban areas where
there was little likelihood of their finding jobs, and back to their home atolls where they could
participate in a more subsistence oriented economy. Namu people living in Majuro in July 1993
agreed that life on their home atoll was better than living in Majuro "because you don't have to
buy food there". For younger people that is not so much a concern as it is for older people with
families.
Another concern is that many of the copra trees on Majkin are old and approaching the end
of their productivity. Unless they are replaced soon there will be a severe shortage of cash for
families to buy the necessary foodstuffs to balance out the times when little or no local foods are
available. Even the Copra Support scheme will be of little use in keeping people on their home
atolls unless the coconut trees are producing enough nuts to be sold to bring in sufficient cash
to feed the increasing population.
Breadfruit trees produce enough fresh fruit to feed the population of Majkin through three
or four months of the year, if eaten at only one meal a day, and supplemented by rice. The
8
fermented paste, bwiro, extends the subsistence base by approximately one month to six weeks,
if eaten only once a day, and if only moderate amounts are given away or sold on Ebeye. As the
number of mouths to be fed increases so these time frames are correspondingly reduced. If
breadfruit is severely hit by cyclone, drought or disease, then this subsistence base is hard
pressed. During 1993 the trees appeared to provide a strong subsistence base. But in another year
the picture may be less rosy. Coconuts are thus crucial as the intermediary between subsistence
and cash food sources. They must be renewed if the population is to continue to maintain at
least a measure of subsistence.
New foods have been added to the inventory since 1968. Pumpkins now are grown
successfully by almost every household and used as an additive to rice, thereby enhancing its
nutritive value. Bananas too have been planted on the borderline between the eolap and iar areas
of each weto. They are eaten as a nutritious snack, in their ripe form. The green banana, widely
used elsewhere in the Pacific is not a familiar source of starch to Marshallese and thus may not
be acceptable, whereas the yellow bananas are. They bring in much desired cash when sold by
the stem on Ebeye.
The planting of taro (iarej) has been encouraged by the Ministry of Resources and
Development in Majuro as an additional starch source. But even though it was growing well on
several weto on Namu, it was not included in the daily diet during 1993. This was partly
because the taste was not wholly acceptable, and partly because it was used formerly as a feast
food only, and not for everyday use. Another drawback was that it takes time, and considerable
fuel to cook. So plants regenerate but are seldom used. However its use may increase with time.
Fuelwood is an increasing problem. At times throughout 1968 fuel for the cooking fires was
in short supply. The people rely mainly on coconut husks for cooking the large pots of rice, or
for roasting the kwonjen form of breadfruit. Such fires are lit at least once a day. As the coconut
trees get older and the number of nuts produced diminishes, so too will the amount of fuel for
cooking fires diminish. Renewal of the coconut trees, plus some attempts to identify suitable
fuelwood trees are two urgent aspects of the subsistence support plan.
The alternative cooking fuel is a kerosene stove. Eleven of the fourteen households had these
in 1993, whereas only four households had them in 1968. This indicates a greater reliance on
kerosene, and thus on cash, for small cooking jobs such as boiling water in a kettle or frying
pancakes. Otherwise the open fire, or the earth oven are used. The kerosene stove is a measure
of a modern lifestyle, as relatives living in Majuro or Ebeye tend to cook on these.
Communication systems have also proliferated. Today every household has one or more
transistor radio, whereas in 1968 only one or two were operating at any one time on the whole
islet, due to shortage of batteries, and/or the radio being broken. As the radio was the only source
of information about field trip ships, it had a major impact on the economy. People relied on
field trip ship itineraries to know when one was coming to Namu so they could make copra. If
they made it too long before the ship arrived, the copra dried out too much and so they lost
money. If they did not have enough time to make copra before the ship arrived they also lost
money.
In addition to the transistor radio, four c.b. aerials are conspicuous additions to the household
9
sites. These are used mainly to talk to friends and relatives in Ebeye and to arrange visits of
people and goods. One of them belongs to Air Marshalls and is used for receiving and sending
information about air traffic.
A weekly air service linking Majkin, Namu to Majuro and Kwajalein, the two urban centres
of the Marshalls, is another major innovation in the communication system. Land on the ocean
side of three weto in the northern part of the islet has been cleared to create a coral runway as
a landing strip. The Air Marshalls 18 seater Dornier lands twice in the same day, once on its way
from Majuro to Namu and then to Ebeye, and once on its return from Ebeye to Majuro via
Namu. Passengers and freight travel regularly, the most heavy traffic being that between Namu
and Ebeye. An additional airstrip exists on Mae-Leuen at the southernmost tip of Namu Atoll.
Majkin people are using the air service as a means of supplying the market for island produce
in the urban centre of Ebeye on Kwajalein Atoll. Boxes of kwonjen, whole pandanus fruits, and
boxes of bwiro, and fish if available, were sent on the plane to Ebeye, either to relatives, or to
be sold. This use of the plane as a means of marketing subsistence produce provides small but
welcome returns to families who have few other alternative sources of cash.
Conclusions
The plant and fish resources of Majkin islet, Namu atoll have continued to be heavily used
over the past twenty five years. The diversity of species, particularly of pandanus, has been
increased to meet specific needs of the handicraft industry, and attempts are being made to
increase the amount of local fish available.
Local resources thus directly provide about 40 per cent of the total local needs today. Copra,
and the cash earned from it indirectly provide another 40 per cent, the cash being spent to buy
the same four basic items as in 1968, rice, flour, tea and sugar. Thus local resources are under
greater pressure today as demands increase. And there are high risks of failure. The balance of
the needs are met by other means, such as support by relatives working in urban centres, wage
labour jobs (though these are still extremely limited) and community support.
The proportion of foodstuffs used locally has remained about the same as that in 1968. But
the possibilities for selling local produce are entirely new, due to new communication systems,
such as c.b. and the airstrip. The pressure on locally produced goods is thus much greater. Two
major differences influence this pattern. Firstly the population of the islet has doubled. It still
maintains its movement patterns from islet to islet and beyond the atoll, so numbers are not
constant; however there are at least twice as many mouths to be fed as in 1968. Secondly the
increased demand for cash has impacted on species other than the coconut for copra. Today
breadfruit, both uncooked and cooked, and its cooked fermented paste as well as pandanus fruits
are highly marketable items among the urban populations. Namu people tend to sell more to
Ebeye because Kwajalein atoll is much closer and communications are better. Some of the goods
are sent by air freight, while other goods are sent in a launch with outboard motor. The cash
realized enables them to buy more rice.
Copra remains as the main source of cash income, but as the trees decline in productivity,
and alternative sources of cash become available, its overall contribution to the economy is
10
declining. There is an urgent need to replace the vast number of older coconut trees, if local
subsistence levels are to be maintained. The population of Majkin "manages" to get by with their
current lifestyle, but would like to have more. Unless rapid and severe measures of population
control are introduced, that lifestyle will not be maintained as there are more mouths to be fed
and educated. More young breadfruit trees also need to be planted as they are slow growing and
will be needed in the future.
The plant resources of Majkin are currently at the interface between subsistence and a cash
economy. More varieties of the same tree crops could be planted, and the coconut trees should
be renewed. These actions would enable the current pattern of about 35 per cent of the food
needs to be met directly from local resources. The other 65 per cent will continue to come from
cash, earned partly from copra, and partly from handicrafts and food sales. By extending
plantings of pandanus and breadfruit further into the middle zone of each weto yields could be
increased. By renewing coconut trees more cash would be available. But so would the risks of
both natural and market disasters.
So whereas local produce in the 1960s was used only for local needs, today that produce is
committed both to local needs and to sales outside the atoll. And with such a rapid increase in
the number of mouths to feed, the atoll is running out of options based on local resources. The
people of Majkin are becoming more and more dependent on outside sources of food, but with
a diminishing supply of cash as copra returns decrease. Highly polished rice is not as nutritious
as breadfruit and pandanus, and it costs money, though it is very popular with the Majkin people.
The people of Majkin face a bleak future. The Marshall Islands government wants to
encourage people to stay on their home islands, and thus reduce the urban pressure. But the
resources on an outer island such as Namu cannot support any further population increase. The
limit has been reached between supply and demand for food. Urgent attention is needed by the
central government to replace coconut trees, increase the increase the number of breadfruit trees,
and to introduce an acceptable means of restricting population size. Otherwise outer island living
will no longer be the idyllic option.
11
BIBLIOGRAPHY
DAMPIER, William, 1697.— A new voyage around the world. Reprint. London 1937.
POLLOCK, Nancy J., 1970. — Breadfruit and Breadwinning on Namu, a Marshallese
atoll. Ph. D thesis, Anthropology. Univ. of Hawaii.
= 1992. — These Roots Remain. Food Habits in islands of the Central and
Eastern Pacific since Western contact. Hawaii: The Institute for
Polynesian Studies.
RAGONE, Diane, 1988. — Breadfruit varieties in the Pacific atolls. UNDP Project
Series. N.Y.: United Nations Development Program.
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ATOLL RESEARCH BULLETIN
NO. 442
CRUSTACEA DECAPODA OF FRENCH POLYNESIA
(ASTACIDEA, PALINURIDEA, ANOMURA, BRACHYURA)
BY
JOSEPH POUPIN
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
APRIL 1996
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FAMILY XANTHIDAE
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CONTENTS
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PAMILY. CRYPTOCHIRIDAE) hiscccecatecaos ces scar ssabecuederennoe cave suse sou cee cue Season ae cea cane thee etek see ee eee 74
PAMIEY LY MENOSOMATIDAE si.s-ccas;csecccesssceusssaurervszeesunacssvsnsdensasese seus soessaduavecessevaeesere corenenemteneee 75
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ICME RAT WIRE! CITED). ccocescclacsuesscessss cuss coors ar cur eas Cae e a eG ee Bee See ee cae eNO Sau ee Rac ee gee 81
ACKNOWLEDGEMENT | cccccctusssccoseccuuicsvocsccoudsstsescssecccnssetecads tsua se usa eee eo ae eee eee eee 95
sas vliesia vai’conuvjiivenosuge douanp oad bGeded exeees sugen ee lebccsu ciate ee aceon MAL: RSI RII CsI Bice ane 95
This work is dedicated to
MONIQUE DALLE
and
JOSETTE SEMBLAT
Librarians at the
Service Mixte de Surveillance Radiologique et Biologique
and Laboratoire de Zoologie des Arthropodes, respectively.
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FRENCH POLYNESIA
CRUSTACEA DECAPODA OF FRENCH POLYNESIA
(ASTACIDEA, PALINURIDEA, ANOMURA, BRACHYURA)
BY
JOSEPH POUPIN
SUMMARY
From a bibliographic compilation and, to a lesser extent, from material collected in the field, 401
littoral and sublittoral decapods (Palinura, Anomura, Brachyura), are reported from French Polynesia. The
Brachyura prevail, with 313 species, mainly Xanthidae (123 species), Portunidae (54 species), and
Grapsidae (35 species). The Anomura are represented by 74 species, and the Palinura by only 14 species.
The list of the deep species, ie living in depths of 100m or more, is updated. Ninety-two species are listed,
making a total of 493 Polynesian species.
Amongst the material recently collected, 16 species are recorded for the first time in the area:
Calcinus guamensis, Calcinus imperialis, Dardanus australis, Dardanus brachyops, Albunea speciosa,
Parthenope contrarius, Portunus macrophthalmus, Portunus orbitosinus, Thalamita danae, Thalamita
macropus, Thalamita mitsiensis, Thalamita philippinensis, Quadrella maculosa, Planes cyaneus, Percnon
guinotae, and Macrophthalmus serenei. Moreover, after the examination of the type material, Ruppelia
granulosa A. Milne Edwards, 1867, originally describe from the Marquesas, is here proposed as a junior
synonym of Lydia annulipes (H. Milne Edwards, 1834). ;
Only 8 species, related to well defined species, are known solely from French Polynesia: Parribacus
holthuisi, Micropagurus polynesiensis, Nucia rosea, Nursia mimetica, Acanthophrys cristimanus,
Lissocarcinus elegans, Ozius tricarinatus, and Macrophthalmus consobrinus. For some of them, however,
it is probable that their distributions extend at least to western Polynesia.
The French Polynesian fauna is typically Indo-West Pacific in its composition, with few endemic
forms, and a low diversity compared to the Indo-Malaysian area. It includes, however, many more species
than the Hawaiian fauna, possibly because the Polynesian islands are less isolated than the Hawaiian
islands.
The Society, Tuamotu, and Gambier archipelagos have been well investigated, with numerous
expeditions organised in these areas. In contrast, the Austral and Marquesas Islands, still remain poorly
known. The French Polynesian fauna is more or less homogenous, with few regionally distinctive features.
The single obvious exception is for the isolated southernmost islands, Rapa and Marotiri, subjected to a
subtropical climate. In these islands, species that are very common elsewhere, are missing (Coenobita
Service Mixte de Surveillance Radiologique et Biologique, SMSRB, B.P. 208, 91311 Montlhéry Cedex,
France, and Muséum national d'Histoire naturelle, Laboratoire de Zoologie des Arthropodes, 61 rue Buffon,
75005 Paris.
Manuscript received 8 December 1995; revised 28 March 1996
perlatus, Birgus latro, Cardisoma carnifex), and, on the contrary, at least one common species is still
unknown in the northern part of French Polynesia (Panulirus pascuensis).
INTRODUCTION
What are the decapod crustacea known from French Polynesia? The answer to this, apparently
simple question, would be very helpful for determinating the species collected during ecological studies.
Moreover, from a biogeographical point of view, a check list of the species reaching this area, at the eastern
limit of the Indo-West Pacific province, would be very interesting. The aim of this work therefore, is to
answer this question by drawing up, mainly from a compilation of systematic and ecological studies, a list
of the French Polynesian crustacea, the scope of the subject being restricted to littoral and sublittoral,
Palinura (Astacidea and Palinuridae), Anomura, and Brachyura.
The check list given here has been mostly compiled from bibliographical records. In a first step, the
most important works dealing with the French Polynesian crustacea have been consulted. They are the
works by DANA (1852b, 1855), HELLER (1865), NOBILI (1907), RATHBUN (1907), BOONE (1934, 1935),
HOLTHUIS (1953), FOREST & GUINOT (1961), and more recently, those by ODINETZ (1983),
MONTEFORTE (1984), GUINOT (1985), MARQUET (1988), and PEYROT-CLAUSADE (1989). In a second
step, the names of the species have been updated, for changes in the generic classification, or species
reduced to synonymy, by looking through more general works, like the ones by HOLTHUIS (1991), SAKAI
(1976), or SERENE (1984). This research has been completed by additional consultation of two
bibliographical journals, the Zoological Records and the Current Contents, and by randomly looking
through reprints available at the Muséum national d'Histoire naturelle, Paris. This last step was sometimes
very fruitful, with additonal species mentioned in the area, often very discreetly.
For the most important families, the main works consulted during this research are the following:
The Palinuridae have been found in the recent catalogue of the Marine lobsters of the world by
HOLTHUIS (1991), and in his revision of the Scyllaridae (HOLTHUIS, 1985).
The pagurids (Coenobitidae, Diogenidae, Paguridae) have been first searched through the work by
NOBILI (1907) and the studies by FOREST, published between 1951 and 1956. Additional information has
been found in the work of LEWINSOHN (1969), the report of RAHAYU (1988), and revisions of the genera,
Aniculus (FOREST, 1984), Calcinus (MORGAN, 1991), Catapaguroides (DE SAINT LAURENT, 1968,
1970), Clibanarius (RAHAYU & FOREST, 1992), Pagurixus (MCLAUGHLIN & HAIG, 1984), and
Trizopagurus (FOREST, 1995).
Except for NOBILI's (1907) work and, for a single species, BOONE's (1935) work, the few
Galatheidae known from French Polynesia come from the ecological works by PEYROT-CLAUSADE
(1977a, b, 1989), KROPP & BIRKELAND (1981), and ODINETZ (1983).
Almost all the Porcellanidae have been found in the works published by HAIG, between 1964 and
1992, HAIG & KROPP (1987), and KROPP (1983, 1986).
For the Brachyura, the beginning of the research has been greatly facilitated by the important studies
of FOREST & GUINOT (1961), MONTEFORTE (1984), and GUINOT (1985). More information has been
found in: MCLAY (1991, 1993), for the Dromiidae; GALIL & CLARK (1994), for the Calappidae of the
genus Matuta; GRIFFIN & TRANTER (1986), for the Majidae; STEPHENSON (1972, 1976), STEPHENSON
& REES (1961, 1967), and MOOSA (1979), for the Portunidae; ODINETZ (1983, 1984a), and the works by
GALIL, and co-authors, published between 1985 and 1990, for the Trapeziidae; SERENE (1984), for the
Xanthidae, and CLARK & GALIL (1993) for the Pilodius xanthids; CROSNIER (1984), for the Carpiliidae
and Menippidae; TURKAY (1973, 1974), for the Gecarcinidae; and SAKAI & TURKAY (1976), CRANE
(1975), and BARNES (1977), for the Ocypodidae of the genera, Ocypode, Uca, and Macrophthalmus,
respectively.
This bibliographical compilation has been completed, in a much more limited way, by the study of
some specimens collected on the field, during the last few years. From the addition of this material, 16
species are recorded for the first time in French Polynesia.
The deep-water crustacea, ie collected from 100m and beyond, have already been listed in POUPIN
(1996), with full references on origins of the collections, and depth ranges. A simple list is produced here,
updated by inclusion of species described after the first compilation, or recently collected (cf. Appendices).
HISTORICAL
OLD VOYAGES: 1820-1900
At that time, the crustacea were collected during the exploring expeditions made around the world by
large sailing vessels. DUPERREY, on board the Coquille (1822-1825), is one of the first to bring back some
species from Tahiti and Bora Bora. They were studied by GUERIN-MENEVILLE (1829, 1838) who
dedicated to DUPERREY a small ocypodid crab from Bora Bora, Gelasimus Duperreyi (now Uca
tetragonon).
About 10 years later (1837-1840), DUMONT D'URVILLE, chief officier of DUPERREY on the
Coquille, sailed again in French Polynesia, commanding the Astrolabe and the Zélée. His vessels visited the
Gambier, Marquesas (Nuku Hiva), and Society Islands. JACQUINOT (1852), naturalist, commanding the
Zélée, mentioned a dozen of species from the area, and described some from the Gambier Islands, including
the small Ocypode pallidula, common on the white sandy beaches of Aukena island.
At the same time, the Americans, worried about participating, like the Europeans, in the discovery of
remote marine areas, launched their first round the world campaign, the great U.S. Exploring Expedition
(1838-1842). The squadron of 6 vessels, commanded by WILKES, left Norfolk in August, 1838. At least
four vessels cruised in French Polynesia: the Flying Fish, Peacock, Porpoise, and the Vincennes. A great
part of the collections from the Tuamotu Islands was lost during the wreck of the Peacock, on the banks of
the Columbia river, however, the Polynesian material, about 10 Anomura and 50 Brachyura, studied by
DANA (1851, 1852a-b, 1855), represents the most important collection from that area. DANA describes
several new species, from Tahiti (Phymodius monticulosus, Trapezia areolata), and the Tuamotu Islands
(Globopilumnus globosus, Liomera tristis, Plagusia speciosa, Thalamita integra, Trapezia bella).
Between 1857 and 1859, the Austrian frigate, Novara, put in at Tahiti, during her sea voyage around
the world. HELLER (1862, 1865) studied the Crustacea of this campaign. He recorded 54 species from
Tahiti, and described several, for example the colourful Calcinus nitidus, and the small gecarcinid,
Epigrapsus politus.
Limited collections were also made at Tahiti by the famous British H.M.S. Challenger (1873-1876).
They are discreetly mentioned in the works of HENDERSON (1888), for the Anomura, MIERS (1886), for
the Brachyura, and BANERJEE (1960), for the grapsid crabs.
This era ends with the voyages of the American ship Albatross (1899-1900, and 1900-1905), and
new collections in the Society, Tuamotu, Gambier, and Marquesas Islands. RATHBUN (1907) studied the
Brachyura collected by this vessel. She recorded 85 Polynesian species and described, for example,
Pachygrapsus fakaravensis, a grapsid very common in the Tuamotu Islands, named after the large atoll of
Fakarava.
French frigate La Coquille at anchor in Matavai bay, Tahiti (1823)
(Drawing by Jules-Louis LEJEUNE. Courtesy of HORIZON Magazine)
BEGINNING OF THE 20TH CENTURY: 1900-1967
The voyages around the world have ended and the collections are now made by people living in
French Polynesia. The most striking in that respect is certainly SEURAT, the head of a small Zoological
laboratory once established at Rikitea, Gambier Islands. Between 1902 and 1905 he gathered an important
collection from the Gambier Islands, and also from the Tuamotu Islands, at Hao and Marutea South. This
material was studied by NOBILI (1906, 1907) with more than 130 Polynesian species, belonging to the
groups here concerned, and with the description of a score of new species, for example Thalamita
gatavakensis, or Thalamita seurati. FOREST (1951), for Calcinus seurati and Calcinus spicatus spp. nov.,
STEPHENSON & REES (1961), for Portunus guinotae sp. nov., and FOREST & GUINOT (1961), in their
study on the Polynesian Brachyura, have mentioned again the material collected by SEURAT.
Gilbert RANSON, of the Malacology department, Muséum national d'Histoire naturelle, Paris, also
collected many specimens. In 1952, during a stay of several months, especially on the atoll of Hikueru, he
gathered numerous scyllarids, pagurids, and crabs. The first two groups have been studied by FOREST
(1953, 1954), with description of 5 new species, including Parribacus holthuisi and Clibanarius ransoni.
The third is studied by FOREST & GUINOT (1961), who, in grouping RANSON and SEURAT material, and
some smaller collections, such as the one made by CHABOUIS, a teacher at the Paul Gauguin school,
Papeete, have registered about 100 crabs, 21 as new records, with some new species, such as Pilumnus
ransoni. The same year, MORRISSON, as a part of the Pacific Sciences Board's Coral Atoll Program,
sampled the most common Crustacea of Raroia atoll, and, to a less extent, of Pukapuka, Takume, and
Tahiti. From that material HOLTHUIS (1953) produced a list of 70 species, and mentioned for the first time
Hippa ovalis, from Tahiti.
During these years, a few expeditions, even if they no longer have the nature of great campaigns
around the world, still visited the Polynesian Islands. For example, SENDLER (1923) recorded about 30
species from Makatea, Rimatara, and Tahiti, from the collections made during the Hanseatischen Siidsee-
Expedition. Some of them, like Coenobita cavipes or the gecarcinid Discoplax longipes, have never been
collected since. In 1931, the yacht Alva explored the Marquesas (Nuku Hiva) and the Society Islands (Bora
Bora, Raiatea, Tahiti). BOONE (1934, 1935) mentioned about 40 species collected during this cruise, and
described two crabs, Actaeomorpha alvae and Lissocarcinus elegans.
Two important expeditions, at an interval of 10 years, mark the end of this period. In 1957, the
Americans organised the Smithsonian Bredin Expedition, which visited the Society and the Tuamotu
Islands. The portunids were studied by STEPHENSON & REES (1967) and STEPHENSON (1976), with
about 30 species, including some new records like Portunus iranjae, Thalamita corrugata, or Thalamita
quadrilobata. GALIL (1985) and, more recently, FOREST (1995), in their works on the genera Tetraloides
and Trizopagurus, respectively, also studied the material of this expedition. In 1967, the boat Pele, during
the Marquesas Expedition, visited the Marquesas, Tuamotu, Society, Gambier Islands, and, in the
neighbourhood, the small island of Pitcairn (HARALD, 1967). The crustacea of this campaign, deposited in
Washington and Perth Museums, have been studied by STEPHENSON (1976), with some fifteen portunids,
and appear, more discreetly, in the studies by SERENE (1972), for Palapedia marquesas sp. nov., GALIL &
LEWINSOHN (1985), for Trapezia tigrina, or HOLTHUIS (1985), for Parribacus holthuisi.
MODERN PERIOD.
In 1966, with the installation of the Centre d’Expérimentation du Pacifique, several scientific
investigations were made, mainly on the atoll of Moruroa. Crustacea collected during these investigations
are mentioned in some systematic works, such as DE SAINT LAURENT (1967), for Catapaguroides fragilis,
or GUINOT (1979) for Lophozozymus superbus, and in ecological studies, for example in CHEVALIER et al.
(1968), SALVAT & RENAUD-MORAND (1969), and LABOUTE & RICHER DE FORGES (1986). These
latter, during the expedition of the old minesweepper Paimpolaise, in the south of French Polynesia
(MacDonald bank), have made the first Polynesian record of Panulirus pascuensis, originally described
from Easter island. From these different campaigns, unstudied collections are still deposited at the Muséum
national d'Histoire naturelle, Paris, for example the crustacea collected by PLESSIS, from which we give
here the first record of Thalamita danae. Others collections were made during the campaigns of the fishing
boat Marara, used by the Direction des Centres d'Expérimentations Nucléaires for monitoring the marine
environment, as a part of the radiological safety program in French Polynesia. Although they mainly
concerned the deep fauna (POUPIN, 1996), some of the most common littoral and sublittoral species were
also collected, and were presented by POUPIN (1994a), in a small illustrated document.
In 1971, the French Ecole Pratique des Hautes Etudes, established its research center in French
Polynesia. First located at Tahiti, it was afterwards transfered to Opunohu Bay, Moorea Island, and is now
denominated Centre de Recherche Insulaire et Observatoire de l'Environnement (CRIOBE). Several
important works have been done by the students or researchers of this center. They are mostly ecological
studies with, however, about 30 new records for the area. The cryptofauna was studied by PEYROT-
CLAUSADE, at Moorea (1977, 1985), and Tikehau (1989). The study of this small fauna was completed by
NAIM (1980) with a dozen of species associated with the algae, at Tiahura, Moorea. The crustacean
associates of the coral Pocillopora, were studied by KROPP & BIRKELAND (1981), and by ODINETZ
(1983, 1984a, b) who described two new Trapezia species, Trapezia serenei and T. punctimanus.
MONTEFORTE (1984) in his Contribution a la connaissance de la faune carcinologique de Polynésie
francaise, collected and studied more than 110 species, some of them, like Calcinus minutus, Calappa
calappa, or Etisus anaglyptus, being new records. More recently, a dozen common species were recorded
from the atoll of Nukutipipi by MERSCHARDT-SALVAT (1991), and the freshwater collections made by
MARQUET (1988, 1991, 1993), have given two new grapsid records, Varuna litterata, from the Society
Islands, and Ptychognathus easteranus, from the Austral and Marquesas Islands.
CONVENTIONS
STUDIES INCLUDED
We have included only studies in which material from French Polynesia has been actually examined,
excluding works where "French Polynesia" appears only in the "Distribution". These are followed by the
indication, in parenthesis, of the island(s) where the material originated. Recent revisions and general
syntheses, from which the names of the species are updated (changes in generic classification; synonymies),
or useful in different aspects, have been added; these are followed by mention such as "Syn." or "Key", in
parenthesis. Some works have been included, that do not mention new collections. These are: the important
syntheses about French Polynesia, like FOREST & GUINOT (1962), followed by "Biogeography", or
GUINOT (1985), followed by "List"; the ecological studies focusing on a particular island, such as
DELESALLE (1985), for the atoll of Mataiva, or SALVAT & RICHARD (1985), for the atoll of Takapoto;
general works dedicated to the fauna of French Polynesia, such as SEURAT (1934), CHABOUIS L. & F.
(1954), or more recently the Encyclopédie de la Polynésie (cf. CHARLEUX, 1986 and SALVAT, 1986a-c)
and the books by PARDON (1992) and BONVALLOT et al. (1994). For these latter, however, only the
species illustrated, usually in colour, have been cited.
No distinction has been made between systematic and ecological works. In her list of the Brachyura
of French Polynesia, GUINOT (1985) has sometimes considered the record of a species doubtful (name
followed by a ?), when it was known only from an ecological paper. As these works can easily be identified
from the references, the reader will be able to judge for himself. It is clear, however, that the revision of
these collections would be important, but it is often difficult, or even impossible, to retrieve the material.
LOCATIONS
The unit of location is the island (see map). When the name of a village, a particular locality, or a
small islet on the recifal crown (Motu in Polynesian language), was indicated, the name of the
corresponding island is mentioned with the following presentation: "Gatavake" = Mangareva, "Ohura" =
Hao, or "Taiohae" = Nuku Hiva. The same presentation is adopted for corrections of obvious mistakes:
"Tickahau" = Tikehau, "Timoe" = Temoe, or "Fakaina" = Fakahina. The Gambier Islands have a particular
configuration, with 8 mountainous islands surrounded by a common recifal crown. In that particular case,
we have considered as real islands three Motu of the external crown: Puaumu, Tarauru-Roa, and
Vaiatekeue.
Sometimes, in the oldest works, the names of the islands were old names, no longer now in use.
They are translated into modern names by using MOTTLER's (1986) work, and the following presentation:
"Carlshoff' = Aratika, "Clermont-Tonnerre" = Reao, or "Eimeo" = Moorea.
When no particular location was specified, the reference is just followed by "French Polynesia".
CLASSIFICATION
The classification approximately follows BOWMAN & ABELE (1982), and, for the Xanthoidea,
SERENE (1984). Subfamily ranks has been indicated only within the most important families: Portunidae,
Xanthidae, and Grapsidae. The presentation has been clarified by ignoring subgeneric names in the check
list. However, if they were used in the works cited, they appear in the references.
LITTORAL, SUBLITTORAL AND DEEP SPECIES
These three groups are here defined in the following way: littoral species are commonly collected on
the reef, and in depth of few meters only; sublittoral species are collected from about 10m to 100m; and
deep species are collected from 100m and deeper. It is sometimes difficult to classify the species according
to these three goups, especially because our knowledge about the deep distribution of numerous species is
often very limited, and will have to be revised in the future. As an example, the maximal depth known for
some species has been increased, here, sometimes considerably, from collections made by traps. Some
littoral forms have been found unusually deep (Carpilius convexus, 60m; Charybdis paucidentata, 100m).
They are qualified as "Littoral to sublittoral" species. Sublittoral forms have been sometimes collected far
beyond 100m (Dardanus brachyops, 110-300m; Dromia wilsoni, 190-350m; Thalamita spinifera, 42-
200m), and, in contrast, deep forms have been found in less than 100m (Palibythus magnificus, 70-240m;
Scyllarus aurora, 90-300m; Alainodaeus rimatara, 90-350m). They are qualified as "Sublittoral to deep"
species. Because of these difficulties, 12 species included in this work were also listed with the deep species
(POUPIN, 1996; see Appendices 1, species with a *).
It is important to realise that, in several cases, these classifications are questionable, and often only
reflect the poor information that we have on that subject. For example, a species like Oreotlos potanus,
known by a single specimen, has been included with the deep species, to within 1m (101m). Considering
the limited accuracy of the measures at sea, O. potanus could have been reasonably considered as a
sublittoral species.
OTHERS CONVENTIONS
When a "?” appears in front of the name of a species, it always means that the doubt on that name is
ours. When it is an hesitation expressed in the work consulted, it is mentioned after the reference by "with a
9"
When a species has been reduced to synonymy, two cases are considered. If the species was not
originally described from French Polynesia, the full name (author and date) appears at the end of the
references, after "SYNONYMS". Otherwise, this information appears clearly with the reference, and is not
repeated again. In both cases the origin of the synonymy is to be found in the work followed by "Syn.". The
synonymies are restricted to French Polynesian species only.
As far as possible we have tried to avoid partial identifications. When a generic name is only
available, the reference is not considered in the main list, but appears separately, in Appendices 2.
Preliminaries identifications (aff. or cf. ) have been retained only when the species refered to is not yet
recorded from French Polynesia (for example, Actaea aff. glandifera in PEYROT-CLAUSADE, 1989: 111).
Otherwise, they appear under the species refered to, after "RELEVANT MATERIAL" (see for example, Lybia
cf. caestifera in MONTEFORTE, 1984: 171, under Lybia caestifera).
The following abbreviations are used: BM (Natural History Museum, London); CRIOBE (Centre de
Recherche Insulaire et Observatoire de l'Environnement, Moorea); MNHN (Muséum national d'Histoire
naturelle, Paris); USNM (National Museum of Natural History, Washington).
LIST OF THE SPECIES
INFRA-ORDER ASTACIDEA
FAMILY ENOPLOMETOPIDAE
Enoplometopus holthuisi Gordon, 1968
DISTRIBUTION. — Tuamotu - Sublittoral.
REFERENCES. — Enoplometopus holthuisi - BONVALLOT et al., 1994: 144-145, photograph (Tuamotu).
REMARK. — At least another Enoplometopus is present in French Polynesia (cf. Enoplometopus sp. nov. in POUPIN ef
al., 1990: 16, pl. 3c).
INFRA-ORDER PALINURIDEA
FAMILY PALINURIDAE
Justitia longimanus (H. Milne Edwards, 1837)
DISTRIBUTION. — Austral (Rurutu); Society (Bora Bora, Raiatea, Tahiti); Tuamotu (Makemo, Maria, Tenarunga) -
Sublittoral to deep.
REFERENCES. — Justitia longimana - POUPIN et al., 1990: 16 (French Polynesia). — Justitia longimanus - POUPIN,
1994b: 46, fig. 3e’, pl. 1d, 2d (Bora Bora, Tahiti, Tenarunga, Rurutu; 62-160m); 1996: in press (Bora Bora, Makemo,
Maria, Raiatea, Rurutu, Tenarunga; 80-190m).
Panulirus homarus (Linné, 1758)
DISTRIBUTION. — Marquesas (Fatu Hiva, Nuku Hiva); Society (Tahit1).
REFERENCES. — Panulirus homarus - GORDON, 1953: 29, fig. 2b-d, 6, 7b (Marquesas "Hana Hevané" = Hanavave
bay?, Fatu Hiva; Puerulus larvae only). — MICHEL, 1971: 467 (Marquesas; Phyllosom larvae only). — HOLTHUIs,
1991: 139, fig. 267-268 (Marquesas, with a ?; Syn.). — ? Panulirus (sic) spinosus (Edwards) - CANO, 1888: 179
(Tahiti) - NEW MATERIAL - Frebruary 1996, Coll. & det. J. POUPIN (Nuku Hiva) - SYNONYMS - Palinurus spinosus H.
Milne Edwards, 1837 (with a ?, in HOLTHUIS, 1991: 139).
Panulirus longipes (A. Milne Edwards, 1868)
DISTRIBUTION. — Marquesas; Society (Tahiti); Tuamotu.
REFERENCES. — Senex femoristriga - ORTMANN, 1891: 23 (Tahiti). — Panulirus longipes - MICHEL, 1971: 467
(Marquesas, Tuamotu; Phyllosom larvae only). — Panulirus longipes femoristriga - HOLTHUIS, 1991: 146, fig. 277b,
278 (Syn.).
REMARK. — In the Indo-West Pacific HOLTHUIS (1991) recognises two subspecies: Panulirus longipes, the western
form, distributed from Africa to Thailand, Taiwan, Indonesia, and Philippines; and P. longipes femoristriga, the eastern
form, known from Japan, the Moluccas, New Guinea, New Caledonia, eastern Australia, and French Polynesia.
Panulirus pascuensis Reed, 1954
DISTRIBUTION. — Austral (MacDonald bank, Marotiri, Rapa) - Littoral to sublittoral.
REFERENCES. — Panulirus pascuensis - LABOUTE & RICHER DE FORGES, 1986: 7, 21, pl. 2c (MacDonald bank,
Marotiri, Rapa; 40m). — SALVAT, 1986b: 70, photograph (MacDonald bank). — HOLTHUIS, 1991: 149, fig. 283-284
(Pitcairn, 500km south-east off the Gambier). — POUPIN, 1994a: 8 (after LABOUTE & RICHER DE FORGES).
REMARK. — LABOUTE & RICHER DE FORGES (1986: 18) also record Panulirus polyphagus (Herbst, 1793) in French
Polynesia ("Iles hautes et atolls"). This species, which is only listed without material examined, is not reported from the
area by HOLTHUIS (1991: 152). We therefore consider that this record is not valid.
Panulirus penicillatus (Olivier, 1791)
DISTRIBUTION. — Gambier; Marquesas; Society (Tahiti); Tuamotu (Makatea, Mataiva, Moruroa, Fakarava, Hao,
Raroia, Taiaro, Takapoto).
REFERENCES. — Panulirus penicillatus - STIMPSON, 1860: 23 [92] (Tahiti). — BATE, 1888: 82, pl. 12-fig. 2 (Tahiti). —
NoBILI, 1907: 366 (Hao). — BOONE, 1935: 67, pl. 17 (Tahiti). — SEURAT, 1934: 60 (Gambier, Tuamotu). — HOLTHUIS,
1953: 50 (Raroia). — CHABOUIS L. & F., 1954: 89 (Tahiti). — MorRIson, 1954: 16 (Raroia). — MICHEL, 1971: 467
(Marquesas, Tuamotu; Phyllosom larvae). — CHEVALIER et al., 1968: 92, 137 (Moruroa). — MONTEFORTE, 1984: 173,
annex 1, tab. a (Makatea, Mataiva, Tahiti, Takapoto). — DELESALLE, 1985: 289, 293 (Mataiva). — SALVAT, 1986b: 70,
photograph (French Polynesia). — BAGNIS & CHRISTIAN, 1983: 108 (Tuamotu). — HOLTHUIS, 1991: 151, fig. 285-286
10
(Tuamotu; Syn.). — PARDON, 1992: 83, photograph (Fakarava). — POUPIN, 1994a: 8, fig. 4 (Taiaro). — Cancer
theresae Curtiss, 1938 (""Tautira" = Tahiti).
Panulirus versicolor (Latreille, 1804)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — ? Panulirus fasciatus - CANO, 1888: 179 (Tahiti; cf. Remark). — Panulirus ornatus - BOONE, 1935:
63, pl. 16 (Tahiti). — POUPIN, 1994a: 8 (French Polynesia; after BOONE, and erroneously after NOBILI, 1907 and
GRUVEL, 1911) - Not Palinurus ornatus (Fabricius, 1798) (cf. Remark). — Panulirus versicolor - HOLTHUIS, 1946:
142, pl. 6-j, pl. 9-b, pl. 11-e,f,m (Tahiti); 1991: 156, fig. 293-294 (French Polynesia).
REMARK. — In HOLTHUIS (1991: 152) Panulirus fasciatus Fabricius, 1798 is a synonym of P. polyphagus (Herbst,
1793). However, CANO's (1888) reference to P. fasciatus would rather be P. versicolor, often recorded under P.
fasciatus (HOLTHUIS, 1991: 152). Moreover, we observe that, in his catalogue, HOLTHUIS (1991) does not mention P.
polyphagus from French Polynesia.
According HOLTHUIS (1946: 140, 142), BOONE's (1935) Tahitian record of Panulirus ornatus (Fabricius, 1798) would
be erroneous, but it is not certain that his material really belongs to P. versicolor.
FAMILY SYNAXIDAE
Palibythus magnificus Davie, 1990
DISTRIBUTION. — Society (Tahiti) - Sublittoral to deep.
REFERENCES. — Palinurellus wienechi (sic) - ANONYMOUS, 1979: 6, 8, 11, not Palinurellus wieneckii (de Man, 1881) =
Palibythus magnificus (Tahiti, 70-240m; material corresponding to the photographs examined and considered by DAVIE
(1990: 686) as "almost definitely of this species"). — Palibythus magnificus Davie, 1990: 686, fig. 1a-b, 3a, c, 4a, 5a
(Tahiti; but not Tuamotu). — POUPIN, 1996: in press (Tahiti, Tuamotu?).
REMARK. — The beautiful set of 10 specimens, collected in 1978 in front of Taravao, Tahiti, has disappeared. It
remains only the photographs examined by DAVIE (1990).
Palinurellus wieneckii (De Man, 1881)
DISTRIBUTION. — Tuamotu - Sublittoral.
REFERENCES. — Palinurellus wieneckii - MICHEL, 1971: 460, fig. 1a-j, tab. 1 (Tuamotu; Puerulus larvae only). —
HOLTHUIS, 1991: 170, fig. 315-316 (Tuamotu: larvae and juveniles; 9-27m).
REMARK. — The larvae, once attributed to this species, could in fact belong to Palibythus magnificus, afterwards
collected in the area (cf. previous species).
FAMILY SCYLLARIDAE
Arctides regalis Holthuis, 1963
DISTRIBUTION. — Tuamotu.
REFERENCES. — Arctides antipodarum - MICHEL, 1971: 467 (Tuamotu; Phyllosom larvae) not A. antipodarum
Holthuis, 1960 = A. regalis, with a doubt, fide HOLTHUIS (1991: 177). — Arctides regalis. — HOLTHUIS, 1991: 177, fig.
331-332 (Tuamotu; Syn.).
Parribacus antarcticus (Lund, 1793)
DISTRIBUTION. — Society (Maupiti, Tahiti); Tuamotu (Anaa, Manihi?, Moruroa, Raroia, Takapoto, Tureia).
REFERENCES. — Scyllarus antarcticus - OWEN, 1839: 86 ("Carysfort" = Tureia). — Parribacus antarcticus - SEURAT,
1934: 60 (Tuamotu). — HOLTHUIS, 1953: 50 (Raroia) pro parte fide HOLTHUIS (1985: 74); 1985: 73, fig. 21, 25a (Anaa,
11
Maupiti, Tahiti, Takapoto; Syn.). — CHABOUIS L. & F., 1954: 89, unnumbered fig. (French Polynesia). — FOREST,
1954b: 345, fig. 26a (Tahiti). — CHEVALIER et al., 1968: 92, 137 (Moruroa). — ? BABLET, 1972: 32, pl. 10 (French
Polynesia). — Parribacus ursus-major - BOONE, 1935: 54, pl. 13 (Tahiti). — Cancer barffi Curtiss, 1938: 164
("Tautira” = Tahiti). — ? "Tiane” - PARDON, 1992: 83, photograph (Manihi) (det. according to the photograph). — Not
Parribacus antarcticus - STIMPSON, 1860: 92 [23] (Tahiti). — NoBILI, 1907: 366 (Hao, "Rikitea” = Mangareva). —
HOLTHUIS, 1953: 50 (Raroia), pro parte. — MorRISON, 1954: 50 (Raroia) - All = Parribacus holthuisi Forest, 1954 fide
HOLTHUIS (1985: 75) - SYNONYMS - Parribacus ursus-major (Herbst, 1793).
Parribacus holthuisi Forest, 1954
DISTRIBUTION. — Gambier (Mangareva); Society (Moorea, Tahiti); Tuamotu (Hao, Hikueru, Mataiva, Raroia,
Takapoto).
REFERENCES. — Parribacus antarcticus - STIMPSON, 1860: 92 [23] (Tahiti). — NOBILI, 1907: 366 (Hao, "Rikitea" =
Mangareva). — HOLTHUIS, 1953: 50 (Raroia) pro parte. — MORRISON, 1954: 50 (Raroia) - All, not P. antarcticus
(Lund, 1793) = P. holthuisi fide HOLTHUIS (1985: 98). — Parribacus holthuisi Forest, 1954b: 346, fig. 25, 26b
(Hikueru, Tahiti). — MONTEFORTE, 1984: 173, annex 1, tab. a (Mataiva, Moorea, Takapoto). — DELESALLE, 1985: 289
(Mataiva). — HOLTHUIs, 1985: 98 (Hao, Hikueru, Mangareva, Tahiti, Raroia). — SALVAT, 1986b: 70, 71, photograph
(French Polynesia).
Parribacus scarlatinus Holthuis, 1960
DISTRIBUTION. — Marquesas (Fatu Hiva).
REFERENCES. — Parribacus scarlatinus - MICHEL, 1971: 472 (Marquesas, Omoa bay = Fatu Hiva). — HOLTHUIS,
1985: 102, fig. 26; 1991: 215, fig. 411-412 (Marquesas).
Scyllarus aurora Holthuis, 1981
DISTRIBUTION. — Austral (Maria, Rurutu, Tubuai); Gambier; Marquesas (Fatu Hiva, Tahuata); Society (Maupiti,
Moorea, Raiatea, Tupai); Tuamotu (Akiaki, Fangataufa, Hao, Makemo, Marutea South, Maria, Moruroa, Tuanake,
Tureia, Vanavana) - Sublittoral to deep.
REFERENCES. — ? Scyllarus sp. TV & V - MICHEL, 1971: 467, tab. 3 (Marquesas, Tuamotu; larvae only). — Scyllarus
aurora Holthuis, 1981: 847, fig. 1-2 (Tubuai; 200m). — MANAC'H & CarsIN, 1985: 473 (Moruroa and/or Fangataufa).
— POUPIN, 1996: in press (Common, 90-300m"; in the distribution, most of the islands are mentioned here for the first
time).
INFRA-ORDER ANOMURA
FAMILY COENOBITIDAE
Birgus latro (Linné, 1767)
DISTRIBUTION. — Gambier (Temoe); Tuamotu (Amanu, Makatea, Marutea South, Matureivavao, Morane, Niau,
Pukapuka, Raroia, Taiaro, Takapoto) - Terrestrial.
REFERENCES. — Birgus latro - DANA, 1852b: 474; 1855, pl. 30, fig. Sa-b (several islands in the Tuamotu; drawing of a
specimen from "Honden" = Pukapuka). — NoBILI, 1907: 375 (Amanu). — SEURAT, 1904a: 242 (Marutea South,
Temoe, "Moture-vavao" = Matureivavao); 1934: 51 (French Polynesia). — SENDLER, 1923: 44 (Makatea). —
HOLTHUIS, 1953: 36 (Raroia). — MOorRISON, 1954: 10 (Raroia). — FOREST, 1954a: 79; 1956a: 1073 (Niau). —
CHABOUIS L. & F., 1954: 92, unnumbered fig. (Makatea). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea,
Takapoto). — DELESALLE, 1985: 288 (Mataiva). — CHARLEUX, 1986: 80, photograph (French Polynesia). — SALVAT &
12
RICHARD, 1985: 356 (Takapoto). — SALVAT, 1986b: 71; 1986c: 8-9, photograph (French Polynesia). — BONVALLOT et
al., 1994: 76, photograph (Tuamotu). — POUPIN, 1994a: 10, fig. 6, pl. 1h (Taiaro) - NEW MATERIAL - Coll. and det. J.
POUPIN (Morane).
REMARK. — GIBSON-HILL (1948: 10) mentions this species from the Marquesas Islands, but it is doubtful that it really
occurs in these Islands, where we have made several unsuccessful investigations.
Coenobita brevimanus Dana, 1852
DISTRIBUTION. — Society (Bora Bora, Tahiti); Tuamotu (Amanu, Hao, Hikueru, Makatea, Mataiva, Niau, Nukutipipi,
Raroia, Takapoto, Taiaro) - Terrestrial.
REFERENCES. — Cenobita clypeata Latr. - HELLER, 1865: 82 (Tahiti). — SEURAT, 1934: 52 (Amanu, Hao). —
Coenobita clypeatus (Herbst) - ORTMANN, 1892a: 316, pl. 12, fig 20 (Tahiti) not C. clypeatus (Herbst, 1794) = C.
hilgendorfi Terao in TERAO (1913: 388). — Coenobita clypeatus Latr. - NOBILI, 1907: 373 (Amanu, "Ohura" = Hao).
— SENDLER, 1923: 42 (Makatea, "Nian" = Niau). — Coenobita hilgendorfi Terao, 1913: 388 (Syn.; cf. Remark). —
FOREST, 1954a: 77 (Hikueru; Syn.); 1956a: 1072 (Hikueru, Tahiti). — Cenobita clypeatus Latreille - BOONE, 1935: 40,
pl. 9 (Bora Bora). — Coenobita brevimanus - HOLTHUIS, 1953: 36 (Raroia). — MORRISON, 1954: 10 (Raroia). —
MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Takapoto). — SALVAT, 1986b: 72 (French Polynesia). —
NAKASONE, 1988: 174 (Syn.). — MERSCHARDT-SALVAT, 1991: 40 (Nukutipipi). — SALVAT F. & B., 1992: 5
(Nukutipipi). — PoOUPIN, 1994a: 11, fig. 7, pl. 1c (Hikueru, Tahiti, Taiaro). — Coenobita ollivieri (sic) - CHARLEUX,
1986: 80-81, photograph (French Polynesia) not C. olivieri (Owen, 1839) = C. brevimanus (correction according to the
photograph).
REMARK. — In TERAO (1913: 389) Coenobita clypeatus (Herbst, 1794) is different from Coenobita clypeatus (Latreille,
1826), and the name Coenobita hilgendorfi is proposed for LATREILLE's material. More recently, NAKASONE (1988)
considers that TERAO's (1913) C. hilgendorfi is the same than C. brevimanus Dana, 1852, and states that, until 1955,
DANA's species has been often referred to as, either C. clypeatus, or C. hilgendorfi.
Coenobita carnescens Dana 1851
DISTRIBUTION. — Tuamotu (Ahe and/or Manihi, Aratika, Kauehi, Pukapuka, Raraka) - Terrestrial.
REFERENCES. — Cenobita carnescens Dana, 1851: 272 (Paumotu); 1852b: 472; 1855, pl. 30, fig. 3a-b ("Carlshoff' =
Aratika, "Honden" = Pukapuka, Raraka, "Vincennes" = Kauehi, "Waterland" = Ahe and/or Manihi). — PouPIN, 1994a:
9, fig. 5 (Text).
REMARK. — In NAKASONE (1988: 165) this species would be valid, although it was considered doubtful by BOUVIER (in
ALCOCK, 1905: 193). According to the drawings provided by DANA, Coenobita carnescens could be in fact the juvenile
form of C. perlatus (cf. POUPIN, 1994a: 12, pl. 1d-f).
Coenobita cavipes Stimpson, 1858
DISTRIBUTION. — Austral (Rimatara) - Terrestrial.
REFERENCES. — Coenobita cavipes - SENDLER, 1923: 43 (Rimatara). — MIYAKE, 1991: 116, fig. 3 (cited only for the
illustration). — POUPIN, 1994a: 9 (Text).
Coenobita olivieri (Owen, 1839)
DISTRIBUTION. — Gambier (Tarauru-Roa); Society (Tahiti) - Terrestrial.
REFERENCES. — Coenobita olivieri - NOBILI, 1907: 374 ("Tarawao, Papenoo" = Tahiti, Tarauru-Roa; cf. Remark). —
SEURAT, 1934: 52 (Tahiti, Gambier). — FOREST, 1956a: 1056 (French Polynesia). — POUPIN, 1994a: 14 (Tahiti; ef.
Remark). — Not Coenobita olivieri - DANA, 1852b: 470 = C. spinosus H. Milne Edwards fide ORTMANN (1892a: 318).
— BAGNIS & CHRISTIAN, 1983: 108, photograph (Tuamotu) = C. perlatus H. Milne Edwards fide POUPIN (1994a: 12).
REMARK. — A specimen attributed by NoBILI to Coenobita olivieri (Owen, 1839) has been examined previously
(PouPIN, 1994a; Tahiti "Papenoo", MNHN Pg2111). It is very close to Coenobita spinosus, and ORTMANN's (1892a)
opinion that Coenobita olivieri is only a variety of C. spinosus, could be justified. Examination of the type material is
required to clarify that point.
13
Coenobita perlatus H. Milne Edwards, 1837
DISTRIBUTION. — Gambier (Mangareva, Tarauru-Roa, Temoe); Society (Moorea, Tahiti); Tuamotu (Amanu, Hao,
Hikueru, Kaukura, Makatea, Marutea South, Mataiva, Moruroa, Nukutipipi, Raroia, Taiaro, Takapoto, Takume) -
Terrestrial.
REFERENCES. — Coenobita perlata - SEURAT, 1904a: 238 (Mangareva, Tarauru-Roa, Temoe, Marutea South); 1904b:
95 (Marutea South); 1934: 51 (French Polynesia). — CHEVALIER ef al., 1968: 85, 137 (Moruroa). — Coenobita
perlatus - NoBILI, 1907: 373 (Amanu, Hao, Kaukura, Marutea, Tarauru-Roa). — SENDLER, 1923: 43 (Makatea,
"Uusuroa, Paumotu" = ?). — HOLTHUIS, 1953: 37 (Raroia). — MORRISON, 1954: 7 (Raroia). — CHABOUIS L. & F.,
1954: 93 ("Mataia" = Tahiti). — FOREST, 1954a: 78; 1956a: 1072 (Hikueru, Takume). — MONTEFORTE, 1984: 172,
annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 6 (Moorea). — DELESALLE, 1985: 288, 289
(Mataiva). — SALVAT & RICHARD, 1985: 359, 360 (Takapoto). — CHARLEUX, 1986: 80-81, photograph (French
Polynesia). — SALVAT, 1986b: 71 (French Polynesia). — MERSCHARDT-SALVAT, 1991: 40 (Nukutipipi). — PARDON,
1992: 83, photograph (Tuamotu). — SALVAT F. & B., 1992: 5 (Nukutipipi). — BONVALLOT et al., 1994: 77, photograph
(Tuamotu). — POUuPIN, 1994a: 12, fig. 8, pl. 1d,f (Hao, Mangareva, Marutea, Taiaro). — Coenobita rugosus vat.
granulatus Bouvier - NOBILI 1907: 373 (Marutea, "Ohura" = Hao) not C. rugosus H. Milne Edwards, 1837 = C.
perlatus fide FOREST (1954a: 78). — Coenobita rugosus - NoBILI, 1907: 373 (Hao, Kaukura) — SEURAT, 1934: 52
(NOBILI's material) - These two references, pro parte not C. rugosus H. Milne Edwards, 1837 = C. perlatus fide FOREST
(1954a: 78). — Coenobita olivieri - BAGNIS & CHRISTIAN, 1983: 108, photograph (Tuamotu) not C. olivieri (Owen,
1839) = C. perlatus fide POUPIN (1994a: 12).
Coenobita rugosus H. Milne Edwards, 1837
DISTRIBUTION. — Austral (Raevavae); Society (Tahiti); Tuamotu (Hikueru, Kaukura, Raraka, Raroia, Takume) -
Terrestrial.
REFERENCES. — Cenobita rugosa - DANA, 1852b: 471; 1855, pl. 30, fig. 1 (Raraka). — STIMPSON, 1858c: 245 [83];
1907: 199 (Tahiti). — HELLER, 1865: 82 (Tahiti). — HENDERSON, 1888: 51 (Tahiti). — SEURAT, 1934: 52 (Kaukura) —
Coenobita rugosus - ORTMANN, 1892a: 317, pl. 12, fig. 22 (Tahiti). — NoBiL, 1907: 373 (Kaukura, "Ohura" = Hao)
pro parte fide FOREST (1954a: 78; some specimens are C. perlatus). — SENDLER, 1923: 42 (Tuamotu). — FOREST,
1954a: 78; 1956a: 1073 (Hikueru, Tahiti, Takume). — HOLTHUIS, 1953: 40 (Raroia). — MORRISON, 1954: 10 (Raroia).
— POUPIN, 1994a: 13, fig. 9, pl. le (Raevavae, Tahiti). — Not C. rugosus (pro parte) and C. rugosus var. granulosa
Bouvier - NoBILI, 1907: 373 = C. perlatus fide FOREST (1954a: 78).
Coenobita spinosus H. Milne Edwards, 1837
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Ahe and/or Manihi, Amanu, Niau, Nukutipipi, Reao) -
Terrestrial.
REFERENCES. — Cenobita olivieri - DANA, 1852b: 470 ("Clermont Tonnerre" = Reao, Tahiti, "Waterland" = Ahe and/or
Manihi) not C. olivieri (Owen, 1839) = C. spinosus fide ORTMANN (1892a: 318). — Coenobita spinosus - NOBILI,
1907: 374 (Amanu). — SENDLER, 1923: 43 ("Nian"=Niau). — SEURAT, 1934: 52 (Amanu). — FOREST, 1956a: 1056
(French Polynesia). — MERSCHARDT-SALVAT, 1991: 40 (Nukutipipi). — SALVAT F. & B., 1992: 5 (Nukutipipi). —
POUPIN, 1994a: 14, fig. 10, pl. 1g (Amanu, Moorea, Tahiti).
FAMILY DIOGENIDAE
Aniculus aniculus (Fabricius, 1787)
DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Moorea, Tahiti); Tuamotu (Ahe and/or
Manihi, Aratika, Fakahina, Hikueru, Makatea, Marokau, Marutea South, Mataiva, Moruroa, Nukutipipi, Rangiroa,
Raraka, Raroia, Takapoto, Tikehau).
REFERENCES. — Aniculus typicus - DANA, 1852b: 461; 1855, pl. 29, fig. 1 ("Carlshoff' = Aratika, Raraka, "Waterland"
= Ahe and/or Manihi). — CANO, 1888: 178 (Tahiti). — Aniculus aniculus - NOBILI, 1907: 371 (Marokau). — SEURAT,
1934: 61(Tuamotu, Gambier). — BOONE, 1935: 36, pl. 8 (Tahiti). — HOLTHUIS, 1953: 41 (Raroia). — MORRISON,
14
1954: 13 (Raroia). — FOREST, 1953b: 561; 1956a: 1067 (Hikueru, Tahiti); 1984: 21, fig. 8, 16, 28-30, 35, 66, 68, 74,
76-85 (Fakahina, Hikueru, Mangareva, Marokau, Marutea South, "Matahiva and Tiahura" = Moorea, Moruroa,
Rangiroa, Raroia, Tahiti, "Tawhae, Marquesas" = Taiohae at Nuku Hiva, "Tikahau atoll"=Tikehau; Syn.). —
MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Moorea, Takapoto); 1987: 8 (Moorea). — DELESALLE,
1985: 289 (Mataiva). — RAHAYU, 1988: 40 (French Polynesia). — MERSCHARDT-SALVAT, 1991: 40 (Nukutipipi). —
SALVAT F. & B., 1992: 5 (Nukutipipi).
REMARK. — Aniculus typicus, proposed by DANA (1852c) in the place of Pagurus aniculus Fabricius, is no more a valid
name (cf. FOREST, 1984: 21).
Aniculus maximus Edmonson, 1952
DISTRIBUTION. — Marquesas (Fatu Hiva, Nuku Hiva).
REFERENCES. — Aniculus maximus - FOREST, 1984: 61, fig. 14, 22, 59-61 (Fatu Hiva, Nuku Hiva). — SALVAT, 1986a:
6, 7, photograph (French Polynesia). — ? Aniculus sp. - SALVAT, 1986b: 71, photograph (French Polynesia; det.
according to the photograph).
Calcinus elegans (H. Milne Edwards, 1836)
DISTRIBUTION. — Gambier (Kamaka, Makaroa, Mangareva); Society (Bora Bora, Tahiti); Tuamotu (Ahe and/or
Manihi, Amanu, Apataki, Aratika, Hao, Hikueru, Marutea South, Mataiva, Raroia, Reao, Tagatau, Taiaro, Takume).
REFERENCES. — Calcinus elegans - DANA, 1852b: 458; 1855, pl. 28, fig. 10a-c (‘Carlshoff' = Aratika, "Clermont
Tonnerre" = Reao, "Waterland" = Ahe and/or Manihi). — HELLER, 1865: 88 (Tahiti). — NOBILI, 1907: 368 (Amanu,
Hao, Kamaka, Makaroa, Marutea South, "Rikitea" = Mangareva, Tagatau, "Wakatihi" = ?). — BOONE, 1935: 23, pl. 3
(Bora Bora, Tahiti). — HOLTHUIS, 1953: 41 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST, 1953b: 555; 1956a:
1062 (Hikueru, Tahiti, Takume). — MONTEFORTE, 1984: 172, annex 1, tab. a (Mataiva). — RAHAYU, 1988: 10, 17
(French Polynesia). — POUPIN, 1994a: 15, fig. 11, pl. 2a (Hao, Taiaro).
Calcinus gaimardi (H. Milne Edwards, 1848)
DISTRIBUTION. — Society (Moorea, Tahiti).
REFERENCES. — Calcinus gaimardi - HELLER, 1865: 87 (Tahiti). — FOREST, 1953b: 555; 1956a: 1062 (Tahiti). —
Kropp & BIRKELAND, 1981: 630, tab. 5 (Moorea). — MONTEFORTE, 1984: 172, annex 1, tab. a; 1987: 8 (Moorea). —
RAHAYU, 1988: 20 (Tahiti).
Calcinus guamensis Wooster, 1984
DISTRIBUTION. — Marquesas (Fatu Hiva, Hiva Oa).
REFERENCES. — Calcinus guamensis - NEW MATERIAL - February 1996, Coll. J. POUPIN, det. J. POUPIN & J. FOREST
(Fatu Hiva, Hiva Oa).
Calcinus imperialis Whitelegge, 1901
DISTRIBUTION. — Society (Tahiti); Tuamotu (Moruroa).
REFERENCES. — Calcinus imperialis - NEW MATERIAL - October 1995, Coll. (in coral Pocillopora) & det. J. POUPIN
(Moruroa, Tahiti "Taravao"). — MORGAN, 1991: 882, fig. 21-23 (Syn.; Key).
Calcinus laevimanus (Randall, 1840)
DISTRIBUTION. — Gambier (Kamaka, Mangareva, Puaumu); Society (Moorea, Tahiti); Tuamotu (Ahe and/or Manihi,
Aratika, Hao, Hikueru, Kauehi, Makatea, Marutea South, Mataiva, Raraka, Raroia, Tagatau, Taiaro, Takapoto,
Takume).
REFERENCES. — Calcinus tibicen - DANA, 1852b: 457 ("Carlshoff' = Aratika, Raraka, "Vincennes" = Kauehi,
"Waterland" = Ahe and/or Manihi) not C. tibicen (Herbst, 1791) = C. laevimanus fide MORGAN (1991: 888). —
HELLER, 1865: 87 (Tahiti; cf. Remark). — HENDERSON, 1888: 61 (Tahiti; of. Remark). — Calcinus herbstii - NOBILI,
1907: 368 pro parte fide FOREST (1951: 84) (Hao, Hikueru, Kamaka, Marutea, "Puamu" = Puaumu, "Rikitea” =
15
Mangareva, Tagatau, Tahiti). — SENDLER, 1923: 42 (Makatea). — FOREST, 1951: 84 (NOBILI's material); 1953: 555;
1956a: 1062 (Hikueru, Tahiti, Takume). — Calcinus herbstii var. lividus Edw. - NoBILi, 1907: 369 (Marutea; cf.
Remark). — Calcinus laevimanus - HOLTHUIS, 1953: 43 (Raroia). — MORRISON, 1954: 7 (Raroia). — MONTEFORTE,
1984: 172, annex 1, tab. a (Makatea, Mataiva, Moorea, Takapoto); 1987: 8 (Moorea). — DELESALLE, 1985: 289
(Mataiva). — RAHAYU, 1988: 10, 18, fig. 1-3 (French Polynesia). — POUPIN, 1994a: 16, fig. 12 (Hikueru, Taiaro) -
SYNONYMS - Calcinus herbstii de Man, 1888; Pagurus lividus H. Milne Edwards, 1848.
REMARK. — Calcinus tibicen (Herbst, 1791) is an Atlantic species. MORGAN (1991: 888) has cited several works in
which C. laevimanus is referred to as C. tibicen, C. herbstii, or C. herbstii var. lividus, but without the works by
HELLER, HENDERSON or NOBILI. Although not verified, this material is here attributed to C. laevimanus.
Calcinus latens (Randall, 1840)
DISTRIBUTION. — Gambier (Vaiatekeue); Society (Moorea, Tahiti); Tuamotu (Fakahina, Hao, Hikueru, Makatea,
Mataiva, Raroia, Taiaro, Takapoto, Takume).
REFERENCES. — Calcinus latens - HELLER, 1865: 88 (Tahiti). — NOBILI, 1907: 369 (Hao). — FOREST, 1951: 84, fig.
14-18 (French Polynesia); 1953b: 556 (Syn.); 1956a: 1062 (Hikueru, Tahiti, Takume). — HOLTHUIS, 1953: 44 (Raroia).
— Morrison, 1954: 7 (Raroia). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — KRopp & BIRKELAND, 1981: 630, tab. 5
(Takapoto). — MONTEFORTE, 1984: 172, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 8 (Moorea). —
GALZIN & POINTIER, 1985: 100 (Moorea). — SALVAT & RICHARD, 1985: 358 (Takapoto). — RAHAYU, 1988: 10, 19
(French Polynesia). — POUPIN, 1994a: 17, fig. 13, pl. 2b (Takume, Taiaro). — Calcinus herbstii - NOBILI, 1907: 368
(Hao, "canal Waiatekene” = Vaiatekeue) pro parte not C. herbstii de Man, synonym of C. laevimanus = C. latens fide
FOREST (1951: 84). — Calcinus terrae-reginae - NOBILI, 1907: 369 (Fakahina, Hao, Mangareva) - SYNONYMS -
Calcinus terrae-reginae Haswell, 1882.
Calcinus minutus Buitendijk, 1937
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Calcinus minutus - MONTEFORTE, 1984: 172, annex 1, tab. a (Moorea).
Calcinus nitidus Heller, 1865
DISTRIBUTION. — Society (Tahiti); Tuamotu (Takapoto).
REFERENCES. — Calcinus nitidus Heller, 1865: 89, pl. 7, fig. 4 (Tahiti). — DE MAN, 1890: 111 (Tahiti). — ORTMANN,
1892a: 293 (Tahiti). — FOREST, 1956b: 218, fig. 14 (Tahiti). — RAHAYU, 1988: 10 (French Polynesia). — POUPIN,
1994a: 18, fig. 14, pl. 2c (Tahiti, Takapoto).
Calcinus seurati Forest, 1951
DISTRIBUTION. — Gambier (Tarauru-Roa, Vaiatekeue); Society (Tahiti); Tuamotu (Hao, Hikueru, Fakahina, Raroia,
Taiaro, Takume).
REFERENCES. — Calcinus herbstii - NOBILI, 1907: 368 (Hao, Fakahina, "Taraourouroa"” = Tarauru-Roa, "chenal
Waiatekene" = Vaiatekeue) pro parte not C. herbstii de Man, 1888, synonym of C. laevimanus = C. seurati fide FOREST
(1951: 86). — Calcinus seurati Forest, 1951: 84, fig. 1, 3-4, 7-8 (NOBILI's material); 1953b: 556; 1956a: 1062
(Hikueru, Tahiti, Takume). — HOLTHUIS, 1953: 44 (Raroia, Takume). — MORRISON, 1954: 7 (Raroia). — RAHAYU,
1988: 19 (Hikueru, Tahiti, Takume). — POUPIN, 1994a: 19, fig. 15, pl. 2d (Hao, Taiaro).
Calcinus spicatus Forest, 1951
DISTRIBUTION. — Gambier (Vaiatekeue).
REFERENCES. — Calcinus herbstii - NOBILI, 1907: 368 ("chenal Waiatekene" = Vaiatekeue) pro parte not C. herbstii de
Man, 1888, synonym of C. laevimanus = C. spicatus fide FOREST (1951: 84). — Calcinus spicatus Forest, 1951: 90,
fig. 10-13 ("chenal Waiatekene" = Vaiatekeue). — RAHAYU, 1988: 21 (French Polynesia).
16
Ciliopagurus krempfi (Forest, 1952)
DISTRIBUTION. — Marquesas (Tahuata) - Sublittoral.
REFERENCES. — Ciliopagurus krempfi - FOREST, 1995: 59 fig. 10c, 11, 12c, 31d, 37g-h (Tahuata; 48m).
Ciliopagurus strigatus (Herbst, 1804)
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Moorea, Tahiti).
REFERENCES. — Pagurus strigatus - ORTMANN, 1892a: 285 (Tahiti). — Trizopagurus strigatus - MONTEFORTE, 1984:
172, annex 1, tab. a (Moorea, Tahiti); 1987: 8 (Moorea). — Ciliopagurus strigatus - FOREST, 1995: 49, fig. 8a, 9, 10a,
12a, 31a-b, 37d (Moorea, Nuku Hiva, Tahiti).
Clibanarius corallinus (H. Milne Edwards, 1848)
DISTRIBUTION. — Society (Bora Bora, Tahiti); Tuamotu (Apataki, Hao, Hikueru, Mataiva, Moruroa, Raroia, Taiaro,
Takapoto, Takume).
REFERENCES. — Clibanarius corallinus - HELLER, 1865: 89 (Tahiti; cf. Remark). — NOBILI, 1907: 367 (Apataki, Hao).
— Boone, 1935: 17, pl. 1 (Bora Bora). — HOLTHUIS, 1953: 45 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST,
1953a: 442; 1956a: 1057 (Hikueru, Tahiti, Takume). — MONTEFORTE, 1984: 172, annex 1, tab. a (Mataiva, Takapoto).
— RAHAYU, 1988: 26, fig. 4-6 (Tahiti). — POUPIN, 1994a: 20, fig. 16 (Apataki, Moruroa, Taiaro).
REMARK. — HELLER (1862: 527) has also described a new Clibanarius from Tahiti, Clibanarius semistriatus.
According to J. FOREST (Personal communication) this species is very doubtful and could be a Pagurus or a Paguristes.
Clibanarius eurysternus Hilgendorf, 1878
DISTRIBUTION. — French Polynesia.
REFERENCES. — Clibanarius eurysternus - RAHAYU, 1988: 10, 28 (French Polynesia). — RAHAYU & FOREST, 1992:
750 (Distribution only "Polynésie").
Clibanarius humilis (Dana, 1851)
DISTRIBUTION. — Gambier (Mangareva, Tarauru-Roa); Society (Tahiti); Tuamotu (Hikueru, Mataiva, Moruroa,
Takume).
REFERENCES. — Clibanarius aequabilis - ? DANA, 1852b: 464; 1855, pl. 29, fig. 4a-f (Tahiti; cf. Remark). — NOBILI,
1907: 367. (""Rikitea" = Mangareva, Tarauru-Roa) not C. aequabilis Dana, 1852 = C. humilis fide FOREST (1953a: 443).
— Clibanarius humilis - FOREST, 1953a: 443, fig. 1, 5; 1956a: 1057 (Hikueru, Tahiti, Takume). — MONTEFORTE,
1984: 172, annex 1, tab. a (Mataiva). — RAHAYU, 1988: 10, 27 (French Polynesia). — POUPIN, 1994a: 21, fig. 17
(Moruroa, Tahiti).
REMARK. — Clibanarius aequabilis Dana, 1852 is an Atlantic species. Concerning DANA's material, FOREST (1953a:
446) writes: "A quoi correspond le sp. C. aequabilis de Tahiti, figuré par Dana ? II] est souhaitable que le terme
d'aequabilis ne soit plus utilisé pour les espéces de ]'Indo-Pacifique."
Clibanarius ransoni Forest, 1953
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Clibanarius ransoni Forest, 1953a: 446, fig. 2, 6; 1956a: 1059 (Tahiti). — RAHAYU, 1988: 30 (Tahiti).
Clibanarius rhabdodactylus Forest, 1953
DISTRIBUTION. — Tuamotu (Hao, Hikueru).
REFERENCES. — Clibanarius zebra - NoBILI, 1907: 367 (Hao), not C. zebra Dana, 1852 = C. rhabdodactylus fide
RAHAYU & FOREST (1992: 777). — Clibanarius zebra var. rhabdodactylus Forest, 1953a: 448, fig. 3, 8; 1956a: 1059
(Hikueru). — RAHAYU, 1988: 29 (French Polynesia). — Clibanarius rhabdodactylus - RAHAYU & FOREST, 1992: 777
(Syn.).
17
Clibanarius striolatus Dana, 1852
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Clibanarius striolatus - HELLER, 1865: 89 (Tahiti). — FOREST, 1953a: 448; 1956a: 1059 (Tahiti). —
RAHAYU, 1988: 10, 27 (Tahiti).
Clibanarius zebra (Dana, 1852)
DISTRIBUTION. — Marquesas (Fatu Hiva, Nuku Hiva, Ua Pou).
REFERENCES. — Clibanarius zebra - FOREST, 1953a: 449, fig. 4 ("Taiohae" = Nuku Hiva) - NEW MATERIAL - Frebruary
1996, Coll. J. POUPIN, det. J. POUPIN & J. FOREST (Fatu Hiva, Nuku Hiva, Ua Pou).
REMARK. — FOREST (1953a) states that the specimens from Marquesas (Coll. Pére Simon DELMAS) are typical of C.
zebra, and distinctly larger than the rhabdodactylus variety.
Dardanus australis Forest & Morgan, 1991
DISTRIBUTION. — Austral (Rapa) - Sublittoral to deep.
REFERENCES. — Dardanus australis - NEW MATERIAL - March 1995, Coll. J. POUPIN, det. J. FOREST (Rapa, 70-115m).
Dardanus brachyops Forest, 1962
DISTRIBUTION. — Marquesas (Tahuata); Society (Bora Bora, Maupiti) - Sublittoral to deep.
REFERENCES. — Dardanus brachyops - POUPIN, 1996: in press (Bora Bora, Maupiti, Tahuata; 110-300m cf. Remark).
REMARK. — This species is still known only beyond 100m in French Polynesia, but it is merely sublittoral in FOREST
(1962: 365; more than 33m to 80m).
Dardanus deformis (H. Milne Edwards, 1836)
DISTRIBUTION. — Gambier (Mangareva?); Society (Tahiti); Tuamotu (Hao, Hikueru, Matureivavao).
REFERENCES. — Pagurus difformis - HELLER, 1865: 86 (Tahiti). — HENDERSON, 1888: 57 (Tahiti). — Pagurus
deformis - NOBILI, 1907: 370 (Hao, and Mangareva with a ?). — BOONE, 1935: 28, pl. 5 (Tahiti). — FOREST, 1953b:
556; 1956a: 1063 (Hikueru, Tahiti). — Dardanus deformis - HOLTHUIS, 1953: 47 (Raroia, Tahiti). — MORRISON, 1954:
7 (Raroia). — RAHAYU, 1988: 35 (French Polynesia) - NEW MATERIAL - Coll. PLESSIS (Matureivavao), coll. C. HILY
(Tahiti), det. J. POUPIN.
Dardanus gemmatus (H. Milne Edwards, 1848)
DISTRIBUTION. — Austral (Maria); Marquesas (Ua Huka); Society (Moorea, Tahiti); Tuamotu (Manihi?, Taiaro) -
Littoral to sublittoral (10-20m).
REFERENCES. — Pagurus gemmatus H. Milne Edwards, 1848: 60 (Marquesas). — STIMPSON, 1858c: 234 [72]
(Marquesas; new material ?). — FOREST, 1953b: 557, fig. 10-11; 1956a: 1063 (Tahiti). — Dardanus gemmatus -
MONTEFORTE, 1984: 172, annex 1, tab. a; 1987: 8 (Moorea). — RAHAYU, 1988: 36 (Tahiti). — POUPIN, 1994a: 22, fig.
18, pl. 2e (Maria, Tahiti, Taiaro, Ua Huka; 10-20m). — Without name - ? SALVAT, 1986a: 23, photograph (Tahiti). —
? PARDON, 1992: 80, photograph (Manihi) - Det. after the photographs.
Dardanus guttatus (Olivier, 1812)
DISTRIBUTION. — Tuamotu (Tureia).
REFERENCES. — Pagurus guttatus - OWEN, 1839: 82 ("Carysfort" = Tureia). — Dardanus guttatus - HAIG & BALL,
1988: 165 (Syn.).
Dardanus lagopodes (Forskal, 1775)
DISTRIBUTION. — Society (Maupiti?, Moorea, Tahiti, Tupai); Tuamotu (Hao, Hikueru, Moruroa).
18
REFERENCES. — Pagurus euopsis - NOBILI, 1907: 370 (Hao). — Pagurus sanguinolentus - FOREST, 1953b: 559, fig.
12-14 (Syn.); 1956a: 1064 (Hikueru, Tahiti). — Dardanus lagopodes - LEWINSOHN, 1969: 32 (Syn.). — MONTEFORTE,
1984: 172, annex 1, tab. a; 1987: 8 (Moorea). — RAHAYU, 1988: 32 (Tahiti, Tuamotu). — "Bernard I'hermite bigaré"
- ? PARDON, 1992: 20, 21, double page photograph (Maupiti; det. according to the photograph) - NEW MATERIAL - Coll.
and det. J. POUPIN (Moruroa, Tahiti, Tupai) - RELEVANT MATERIAL - Dardanus aff. sanguinolentus - SALVAT &
RENAUD-MOoRNANT, 1969: 165 (Moruroa) - SYNONYMS - Pagurus euopsis Dana, 1852; P. sanguinolentus Quoy &
Gaimard, 1824.
Dardanus megistos (Herbst, 1804)
DISTRIBUTION. — Society (Tahiti); Tuamotu (Hao, Raroia, Tagatau, Taiaro).
REFERENCES. — Pagurus spinimanus Edw. - DANA, 1852b: 452 (with a ?); 1855, pl. 28, fig. Sa-c (Tuamotu). —
BOONE, 1935: 34, pl. 7 (Tahiti). — Pagurus punctulatus Olivier - HELLER, 1865: 87 (Tahiti). — NoBILI, 1907: 370
("Ohura" = Hao, Tagatau). — Pagurus megistos - FOREST, 1953b: 559; 1956a: 1064 (Tahiti). — Dardanus megistos -
HOLTHUIS, 1953: 49 (Raroia). — MORRISON, 1954: 7 (Raroia). — RAHAYU, 1988: 33 (French Polynesia). — POUPIN,
1994a: 23, fig. 19, pl. 2g (Tahiti, Taiaro). — Without name - ? SALVAT & RIVES, 1975: 57, full page photograph
(French Polynesia; det. according to the photograph) - SYNONYMS - Pagurus punctulatus Olivier, 1811; P. spinimanus
H. Milne Edwards, 1848.
Dardanus pedunculatus (Herbst, 1804)
DISTRIBUTION. — Austral (Tubuai); Tuamotu (Moruroa) - Littoral to sublittoral.
REFERENCES. — Dardanus haani - CHEVALIER et al., 1968: 119 (Moruroa). — SALVAT & RENAUD-MORNANT, 1969:
165, 176 (Moruroa). — Dardanus pedunculatus - RAHAYU, 1988: 10, 34 (French Polynesia). — POUPIN, 1994a: 24,
fig. 20, pl. 2f (Tubuai; 65m) - SYNONYMS - Dardanus haani (Rathbun, 1902).
Dardanus scutellatus (H. Milne Edwards, 1848)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Pagurus scutellatus - FOREST, 1953b: 560; 1956a: 1066 (Tahiti). — Dardanus scutellatus - RAHAYU,
1988: 36 (Tahiti).
Diogenes gardineri Alcock, 1905
DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Marutea South).
REFERENCES. — Diogenes gardineri - NOBILI, 1907: 366 (Marutea South; "Rikitea" = Mangareva). — SEURAT, 1934:
61 (Mangareva, Marutea South). — FOREST, 1956a: 1056 (French Polynesia); 1957: 530 (NOBILI's material).
FAMILY PAGURIDAE
Catapaguroides fragilis (Melin, 1939)
DISTRIBUTION. — Tuamotu (Moruroa) - Littoral to sublittoral.
REFERENCES. — Catapaguroides fragilis - DE SAINT LAURENT, 1968: 940, fig. 26 with hesitations (cf. p. 941, note 1)
(Moruroa; 40m). — CHEVALIER et al., 1968: 119, 138 (Moruroa).
Micropagurus polynesiensis (Nobili, 1906)
DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Moruroa).
REFERENCES. — Anapagurus polynesiensis Nobili, 1906a: 260; 1907: 372, pl. 1, fig. 10. ("Rikitea" = Mangareva). —
FOREST, 1956a: 1056 (French Polynesia). — CHEVALIER ef al., 1968: 119 (Moruroa). — Micropagurus polynesiensis -
HAIG & BALL, 1988: 184 (Syn.).
19
Pagurixus anceps (Forest, 1954)
DISTRIBUTION. — Society (Tahiti); Tuamotu (Hikueru).
REFERENCES. — Eupagurus anceps Forest, 1954a: 71, fig. 15-19; 1956a: 1067 (Hikueru, Tahiti). — ELDREDGE, 1967:
13 (Hikueru). — Pagurixus anceps - MCLAUGHLIN & HAIG, 1984: 135, fig. 5 (Hikueru; Syn.).
Pagurixus laevimanus (Ortmann, 1892)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Eupagurus laevimanus Ortmann, 1892a: 302, pl. 12, fig. 13 (Tahiti). — Pagurixus laevimanus -
MCLAUGHLIN & HAIG, 1984: 142, fig. 7 (Syn.).
Pagurixus maorus (Nobili, 1906)
DISTRIBUTION. — Gambier (Mangareva).
REFERENCES. — Eupagurus maorus Nobili, 1906a: 259; 1907: 371, pl. 1, fig. 9 (Mangareva). — FOREST, 1954a: 73;
1956a: 1056 (French Polynesia). — Pagurixus maorus - MCLAUGHLIN & HAIG, 1984: 126, fig. 2 (Syn.). — KOMAI &
ASAKURA, 1995: 341, 353 (Key).
Trichopagurus trichophthalmus (Forest, 1954)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Catapaguroides trichophthalmus Forest, 1954a: 74, fig. 20-24, with a ? for the genus; 1956a: 1069
(Tahiti). — Trichopagurus trichophthalmus - DE SAINT LAURENT, 1970: 212, fig. 1-16 (Tahiti).
FAMILY GALATHEIDAE
Coralliogalathea humilis (Nobili, 1905)
DISTRIBUTION. — Society (Moorea); Tuamotu (Hao, Tikehau).
REFERENCES. — Galathea megalochira Nobili, 1906a: 260; 1907: 376, pl. 1, fig. 12 (Hao). — Coralliogalathea
humilis - LEWINSOHN, 1969: 117 (Syn.). — PEYROT-CLAUSADE, 1977a, annex of the species: 24 (Moorea); 1989: 113,
115 (Moorea, Tikehau). — KRopp & BIRKELAND, 1981: 629, tab. 5 (Moorea). — ? Galathea himilis (sic) - PEYROT-
CLAUSADE, 1977b: 213 (Moorea).
Galathea aculeata Haswell, 1882
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Takapoto).
REFERENCES. — Galathea aculeata - ODINETZ, 1983: 208 (Moorea, Tahiti, Takapoto). — ODINETZ-COLLART & RICHER
DE ForGES, 1985: 201 (Moorea and/or Tahiti, Takapoto). — MONTEFORTE, 1987: 8 (Moorea).
Galathea aff. amamiensis Miyake & Baba, 1966
DISTRIBUTION. — Society (Moorea) - Littoral to sublittoral.
REFERENCES. — Galathea aff. amamiensis - PEYROT-CLAUSADE, 1989: 115 (Moorea; 30m).
Galathea affinis Ortmann, 1892
DISTRIBUTION. — Gambier (Mangareva); Society (Moorea); Tuamotu (Hao, Marutea South, Tikehau) - Littoral to
sublittoral.
REFERENCES. — Galathea affinis - NOBILI, 1907: 375, pl. 1, fig. 11 (Marutea South, "Ohura" = Hao, "Rikitea and
Gatavake" = Mangareva). — PEYROT-CLAUSADE, 1977a, annex of the species: 24; 1977b: 213; 1985: 462 (Moorea);
1989: 112, 115 (Moorea, Tikehau; 30m). — KRopp & BIRKELAND, 1981: 630, tab. 5 (Moorea).
20
Galathea algae Baba, 1969
DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral.
REFERENCES. — Galathea algae - PEYROT-CLAUSADE, 1989: 112, 115 (Moorea, Tikehau; 30m).
Galathea latirostris Dana, 1852
DISTRIBUTION. — Society (Raiatea, Tahiti).
REFERENCES. — Galathea latirostris - BOONE, 1935: 50, pl. 12 (Raiatea, Tahiti).
REMARK. — The status of this species, originally described by DANA (1852b: 480; 1855, pl. 30, fig. 8) from the Fiji is
doubtful. The examination of topotypic material, and the revision of all the references attributed to Galathea latirostris
is necessary to establish its real identity (K. BABA, personal communication). Galathea latirostris Lenz, 1902, describes
form Juan Fernandez must be attributed to Phylladiorhynchus pusillus (Henderson, 1885) (cf. BABA, 1991: 487), and
must not be confounded with DANA's species.
Phylladiorhynchus serrirostris (Melin, 1939)
DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral.
REFERENCES. — Galathea serrirostris - PEYROT-CLAUSADE, 1977b: 213 (Moorea). — Phylladiorhynchus serrirostris -
PEYROT-CLAUSADE, 1977a, annex of the species: 24 (Moorea); 1989: 112, 115 (Moorea, Tikehau; "...extremely
abundant at 30m"). — KRopP & BIRKELAND, 1981: 630, tab. 5 (Moorea).
Sadayoshia miyakei Baba, 1969
DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral.
REFERENCES. — Sadayoshia miyakei - KROPP & BIRKELAND, 1981: 630, tab. 5 (Moorea). — PEYROT-CLAUSADE, 1989:
112, 115 (Moorea, Tikehau; 30m).
FAMILY PORCELLANIDAE
Neopetrolisthes maculatus (H. Milne Edwards, 1837)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Neopetrolisthes oshimai (sic) - PARDON, 1992: 81 (Tahiti). — Neopetrolisthes maculatus - HAIG,
1979: 127 (Syn.).
REMARK. — According to HAIG (1979), Neopetrolisthes ohshimai Miyake, 1937 is a synonym of this species. This
small crustacea is commensal of a sea-anemone. PARDON (1992) illustrates a shrimp Stenopus captured in a sea-
anemone. "Neopetrolisthes oshimai" is only indicated in the caption, but is not visible on the photograph.
Pachycheles pisoides (Heller, 1865)
DISTRIBUTION. — Society; Tuamotu (Tikehau).
REFERENCES. — Pachycheles pisoides - HAIG, 1966: 290 (Tuamotu; with the mention that it is the first record for the
area, but without details on the material examined); 1983: 284 (Distribution only "Society Islands"). — PEYROT-
CLAUSADE, 1989: 113 (Tikehau).
Pachycheles sculptus (H. Milne Edwards, 1837)
DISTRIBUTION. — Society (Moorea); Tuamotu.
REFERENCES. — Pachycheles sculptus - HAIG, 1966: 287 (Tuamotu; same remark than for the previous species); 1983:
284 (Distribution only "Tuamotu"); 1992: 310 (Syn.), — PEYROT-CLAUSADE, 1989: 115 (Moorea).
Petrolisthes bispinosus Borradaile, 1900
DISTRIBUTION. — French Polynesia.
REFERENCES. — Pefrolisthes bispinosus - HAIG & KRoppP, 1987: 171, 172, fig. 1-2 (French Polynesia, only in summary
and discussion).
Petrolisthes borradailei Kropp, 1983
DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Huahine, Moorea, Tahiti); Tuamotu
(Fakarava, Makemo, Moruroa, Rangiroa, Raroia, Taiaro, Tikehau).
REFERENCES. — Petrolisthes rufescens - NOBILI, 1907: 377 ("Rikitea" = Mangareva; cf. Remark under P. rufescens).
— PoupPIN, 1994a: 25, fig. 21, pl. 2h (Mangareva, Moruroa, Taiaro) not P. rufescens = P. borradailei fide KROPP
personal communication. — Petrolisthes borradailei Kropp, 1983: 96, 106, fig. 3 ("Fakarova" = Fakarava, Huahine,
"Makeno" = Makemo, Moorea, Nuku Hiva, Rangiroa, Raroia, "Pascua Pass" = ?, Tahiti, "Tikahua" = Tikehau).
Petrolisthes coccineus (Owen, 1839)
DISTRIBUTION. — Society (Moorea); Tuamotu.
REFERENCES. — Porcellana coccinea Owen, 1839: 87, pl. 26, fig. 1-2 ("Low Islands" = Tuamotu). — Petrolisthes
coccineus - PEYROT-CLAUSADE, 1977a, annex of the species: 24; 1977b: 213 (Moorea). — HAIG, 1983: 280; 1992: 313,
fig. 9 (Syn.).
Petrolisthes decacanthus Ortmann, 1897
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Petrolisthes decacanthus Ortmann, 1897a: 285, pl. 17, fig. 2 (Tahiti). — Haic & KRopp, 1987: 176
(French Polynesia, in the distribution only; Syn.).
Petrolisthes eldredgei Haig & Kropp, 1987
DISTRIBUTION. — Society (Tahiti); Tuamotu (Raroia).
REFERENCES. — Petrolisthes eldredgei Haig & Kropp, 1987: 180, fig. 5-6 (Tahiti, Raroia).
Petrolisthes elegans Haig, 1981
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava).
REFERENCES. — Petrolisthes bosci - NoBILI, 1907: 377 ("“Sakarava" = Fakarava) not P. bosci (Audouin, 1826) = P.
elegans sp. nov. in HAIG (1981: 266). — Petrolisthes elegans Haig, 1981: 266, fig. 2 (Tahiti, "Sakarava" = Fakarava).
— PEYROT-CLAUSADE, 1989: 115 (Moorea).
Petrolisthes lamarckii (Leach, 1820)
DISTRIBUTION. — Tuamotu (Raraka, Takaroa).
REFERENCES. — Porcellana speciosa Dana, 1852b: 417; 1855, pl. 26, fig. 8 (Raraka). — EVANS, 1967: 409 (Raraka;
syntypes at the BM). — Petrolisthes lamarckii - HAIG, 1964: 362 (Takaroa); 1992: 315, fig. 11 (Syn.). — KRopp, 1983:
100, 106 (Syn.).
Petrolisthes militaris (Heller, 1862)
DISTRIBUTION. — Tuamotu (Marutea South).
REFERENCES. — Petrolisthes militaris - NOBILI, 1907: 377, with a ? (Marutea South).
REMARK. — The presence of this species in French Polynesia is doubtful. NOBILI's determination is uncertain and,
moreover, it has never been reported in the area by HAIG (1979: 122; 1982: 280; 1992: 316).
22
Petrolisthes pubescens Stimpson, 1858
DISTRIBUTION. — Marquesas (Nuku Hiva).
REFERENCES. — Pefrolisthes pubescens - KRopP, 1986: 456, fig. 2 ("Taiohae, Haka Paa, baie du Controleur" = Nuku
Hiva).
Petrolisthes rufescens (Heller, 1861)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Porcellana rufescens - HELLER, 1865: 76 (Tahiti). — Not Petrolisthes rufescens - NOBILI, 1907: 377
(cf. Remark). — PouPIN, 1994a: 25, fig. 21, pl. 2h = P. borradailei Kropp, 1983 fide KROpP personal communication.
REMARK. — Petrolisthes rufescens and P. borradailei are two very close species. The main difference concerned the
posterior border of the cheliped carpus, strongly toothed in P. borradailei, more smooth in P. rufescens. We have
confused these two species in a previous work (POUPIN, 1994a) and re-examination of NOBILI's material in Paris (9 ov.
8x7.3, MNHN Ga96) show that it also belongs to P. borradailei. HELLER's reference remains the only record of P.
rufescens in the pacific and could also belongs to P. borradailei.
Petrolisthes scabriculus (Dana, 1852)
DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral.
REFERENCES. — Petrolisthes scabriculus - PEYROT-CLAUSADE, 1977a, annex of the species: 25; 1977b: 213 (Moorea);
1989: 112, 115 (Moorea, Tikehau; 30m). — HAIG, 1979: 120 (Syn.).
Petrolisthes tomentosus (Dana, 1852)
DISTRIBUTION. — Society (Tahiti); Tuamotu (Raraka).
REFERENCES. — Porcellana tomentosa Dana, 1852b: 420; 1855, pl. 26, fig. 10 (Raraka). — Petrolisthes tomentosus -
Kropp, 1986: 453, fig. 1 (Tahiti, Tahitian neotype at the USNM; Syn.).
Porcellana mitra Dana, 1852
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Porcellana mitra - HELLER, 1865: 74, 265, ("Siidsee" and Tahiti; cf. Remark).
REMARK. — In HELLER, Tahiti is not mentioned in the main text (p. 74), but only at the end of the work, in the part
concerning the geographical distribution (p. 265). It is thus not certain that the location "Tahiti" is correct for this
species.
Porcellana monilifera Dana, 1852
DISTRIBUTION. — Tuamotu (Raraka?).
REFERENCES. — Porcellana monilifera Dana, 1852b: 413; 1855, pl. 26, fig. 3 (Raraka, with a ?).
FAMILY ALBUNEIDAE
Albunea speciosa Dana, 1852
DISTRIBUTION. — Society (Moorea?).
REFERENCES. — Albunea speciosa - NEW MATERIAL - CRIOBE collections, Moorea (without label), 1 d 11x10.5, det.
J. POUPIN.
REMARK. — This specimen agrees very well with DANA's species, illustrated by SERENE (1973). Formely, Albunea
speciosa was considered as endemic from Hawaii. SERENE (1973) has examined specimens from the type locality but
mentions that the type material has disappeared. In 1973, THOMASSIN, has described A. madagascariensis, very close to
A. speciosa. By the shape of the ocular peduncle and the number of frontal spines, the specimen from Moorea do
belongs to DANA's species.
FAMILY HIPPIDAE
Hippa adactyla Fabricius, 1787
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti) - Brackish water.
REFERENCES. — Remipes testudinarius - ? HELLER, 1865: 72 (Tahiti; reference not found in HAIG, 1970). — DE Man,
1896: 466 ("Nuka-hiwa (Mus. Paris)" = Nuku Hiva). — Hippa adactyla - HOLTHUIS, 1953: 35 (Tahiti). — HAI, 1970:
294 (Syn.); 1974: 179, fig. 2, pl. 6 (Marquesas, distribution only) - NEW MATERIAL - 1 6 28x23.5, coll. C. HILY, det. J.
POUPIN ("‘Arue" = Tahiti; confronted with DE MAN's material) - SYNONYMS - Remipes testudinarius Latreille, 1806.
Hippa ovalis (A. Milne Edwards, 1863)
DISTRIBUTION. — Society (Tahiti) - Brackish water.
REFERENCES. — Hippa ovalis - HOLTHUIS, 1953: 35 (Tahiti). — THOMASSIN, 1969: 154, fig. 7b, 8b, 9, pl. 6, fig. 1-8
(Syn.).
Hippa pacifica (Dana, 1852)
DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti) - Brackish water.
REFERENCES. — Remipes pacificus - NOBILI, 1907: 378 (Mangareva). — SEURAT, 1934: 60 (Tahiti). — ? Hippa sp. -
CHABOUIS L. & F., 1954: 92, unnumbered fig. (French Polynesia; det. according to the shape of the front margin, on the
figure). — Hippa pacifica - HAIG, 1974: 181, fig. 3 (Gambier, distribution only; Syn.).
REMARK. — NOBILI's material has been re-examined (MNHN Hi38, Mangareva, 1 2 ov. 22x17, coll. SEURAT 1905,
det. NOBILI 1906) and is similar to other specimens attributed by DE MAN to this species (MNHN Hi107, Java sea;
MNHN Hi108, Atjeh).
INFRA-ORDER BRACHYURA
FAMILY DROMIIDAE
Cryptodromia coronata Stimpson, 1859
DISTRIBUTION. — Gambier (Mangareva).
REFERENCES. — Cryptodromia coronata - NoBILI, 1907: 378 ("Rikitea" = Mangareva). — FOREST & GUINOT, 1962: 56
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 448 (List). — Cryptodromia ? coronata - MCLAY, 1993: 199
(Syn.; with the indication that the status of the species is uncertain).
Cryptodromia fallax (Lamarck, 1818)
DISTRIBUTION. — Tuamotu (Raroia).
REFERENCES. — Cryptodromia canaliculata - HOLTHUIS, 1953: 3 (Raroia). — MORRISON, 1954: 13 (Raroia). —
FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 448 (List). — Cryptodromia fallax -
McLay, 1993: 206, fig. 18e (Syn.) - SYNONYMS - Cryptodromia canaliculata Stimpson, 1858.
24
Cryptodromiopsis tridens Borradaile, 1903
DISTRIBUTION. — Society (Moorea, Tahiti).
REFERENCES. — Cryptodromiopsis tridens - MCLAY, 1991: 467, fig. 5a-d (Moorea, Tahiti).
Dromia wilsoni (Fulton & Grant, 1902)
DISTRIBUTION. — Austral (Raevavae); Marquesas (Tahuata); Tuamotu (Makemo, Takapoto) - Sublittoral to deep.
REFERENCES. — Petalomera wilsoni - MCLAY, 1991: 470, fig. 6a-d, 7a-c, 8a-c (Makemo, Raevavae, Tahuata,
Takapoto; 190-350m). — Dromia wilsoni - MCLAY, 1993: 156, fig. 16e (Syn.). — POUPIN, 1996: in press (same
material than MCLAY).
REMARK. — Dromia wilsoni is mainly a sublittoral species, usually found within the first 100m, but it is also recorded
up to 520m (cf. in McLay, 1991: 475).
FAMILY DYNOMENIDAE
Dynomene hispida Desmaret, 1825
DISTRIBUTION. — Society (Moorea); Tuamotu (Marutea South, Tikehau).
REFERENCES. — Dynomene hispida - NoBILI, 1907: 378 (“Marutea-Vaitutaki" = Marutea South). — FOREST & GUINOT,
1962: 56 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 25; 1977b: 212
(Moorea); 1989: 111, 114 (Moorea, Tikehau). — GUINOT, 1985: 448 (List).
Dynomene praedator A. Milne Edwards, 1879
DISTRIBUTION. — Society (Moorea, Tahiti).
REFERENCES. — Dynomene praedator - ORTMANN, 1892b: 534, pl. 26, fig. 3 (Tahiti). — FOREST & GUINOT, 1962: 56
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 448 (List). — Dynomene sp. - NAIM, 1980a: 55, fide observation
and personal communication of MCLAY (Moorea, MNHN B20203). — Dynomene sinense - ODINETZ, 1983: 208
(Tahiti; MNHN B17090). — Dynomene sinensis (sic) - GUINOT, 1985: 448 (List; after ODINETZ) - These two
references, not D. sinense Chen, 1979 = D. praedator fide observation and personal communication of MCLAY.
Dynomene spinosa Rathbun, 1911
DISTRIBUTION. — Marquesas; Tuamotu (Raroia).
REFERENCES. — Dynomene spinosa - BALSS, 1935: 115 (Marquesas). — HOLTHUIS, 1953: 3 (Raroia). — MORRISON,
1954: 13 (Raroia). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"; Marquesas). — GUINOT, 1985:
448 (List).
FAMILY RANINIDAE
Notosceles chimmonis Bourne, 1922
DISTRIBUTION. — Marquesas (Eiao) - Sublittoral to deep.
REFERENCES. — Notosceles chimmonis - POUPIN, 1996: in press (Eiao; 54-101m).
REMARK. — Species known between 45-52 m (SERENE & UMALI, 1972), 75-90m (RIBES, 1989), and up to 450m
(Monop, 1975).
Ranina ranina (Linné, 1758)
DISTRIBUTION. — Marquesas (Eiao, Fatu Hiva, Nuku Hiva) - Sublittoral.
25
REFERENCES. — Ranina ranina - GUINOT, 1985: 449 (List; certainly after the following dry specimen: MNHN n°223
"Nouhiva" = Nuku Hiva) - NEW MATERIAL - Coll. and det. J. POUPIN (Eiao, Fatu Hiva; juveniles, 100m).
FAMILY CALAPPIDAE
Ashtoret lunaris (Forskal, 1775)
DISTRIBUTION. — Marquesas (Nuku Hiva).
REFERENCES. — Matuta banksii - RATHBUN, 1907: 68 (Nuku Hiva). — GUINOT, 1985: 453 (List). — Ashtoret lunaris -
GALIL & CLARK, 1994: 5, fig. 1a-b, pl. la-b (Syn.) - SYNONYMS - Matuta banksi Leach, 1817.
Ashtoret picta (Hess, 1865)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Matuta picta - POUPIN, 1994a: 27, fig. 23, pl. 3b (Tahiti). — Ashtoret picta - GALIL & CLARK, 1994:
18, fig. 3c-d, pl. 6a-b (Tahiti).
REMARK. — In the revision of the genus Matuta by GALIL & CLARK (1994), the location "Tahiti", that should appear
under Ashtoret picta, is erroneously mentioned under Ashtoret granulosa (Miers, 1877), species still unknown from
French Polynesia (GALIL, personal communication).
Calappa calappa (Linné, 1758)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Calappa calappa - MONTEFORTE, 1984: 173, annex 1, tab. a (Tahiti; MONTEFORTE's material verified
in the CRIOBE collections, Moorea). — GUINOT, 1985: 449 (List).
Calappa hepatica (Linné, 1758)
DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Tahiti); Tuamotu
(Hao, Hikueru, Marutea North?, Marutea South, Mataiva, Moruroa, Takapoto).
REFERENCES. — Calappa hepatica - NoOBILI, 1907: 378 ("Gatavake" = Mangareva, Hao, Marutea = Marutea North?,
Marutea South). — RATHBUN, 1907: 67 (Bora Bora). — PESTA, 1913: 37 (Tahiti). — BOONE, 1934: 32, pl. 8-10 (Nuku
Hiva). — SEURAT, 1934: 59, 60 (Hao, Marutea South). — CHABOUIS L. & F., 1954: 92, unnumbered fig. (French
Polynesia). — FOREST & GUINOT, 1961: 11, fig. 1a-b, 2 (Hikueru; Syn.); 1962: 56 (Biogeography "Tahiti-Tuamotu;
Marquesas"). — MONTEFORTE, 1984: 173, annex 1, tab. a, photograph p. 140c (Mataiva, Moorea, Tahiti, Takapoto);
1987: 8 (Moorea). — DELESALLE, 1985: 288 (Mataiva). — GUINOT, 1985: 449 (List). — POUPIN, 1994a: 26, fig. 22, pl.
3a (Hikueru, Moruroa, Tahiti). — Calappa tuberculata - HELLER, 1865: 69 (Tahiti) - SYNONYMS - Calappa tuberculata
Fabricius, 1798.
Matuta victor (Fabricius, 1781)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Matuta victor - HELLER, 1865: 69 (Tahiti). — GALIL & CLARK, 1994: 39, fig. 7a-b, pl. 13a-b (cf.
Remark).
REMARK. — HELLER's reference is not mentioned in the revision by GALIL & CLARK (1994). These authors consider
that the eastern distribution of this species is limited to the Fiji. Thus, the revision of HELLER's material would be
important to confirm the presence of Matuta victor in French Polynesia.
26
FAMILY LEUCOSIIDAE
Ebaliopsis erosa (A. Milne Edwards, 1874)
DISTRIBUTION. — Gambier; Tuamotu (Marutea South).
REFERENCES. — Ebalia erosa - NOBILI, 1907: 378 (Marutea). — Ebaliopsis erosa - FOREST & GUINOT, 1962: 56
(Biogeography "Tahiti-Tuamotu"). — SERENE, 1977: 55, fig. 5-6 (Gambier). — GUINOT, 1985: 453 (List).
Heteronucia venusta Nobili, 1906
DISTRIBUTION. — Society (Moorea); Tuamotu (Fakahina, Fakarava, Hao, Tikehau).
REFERENCES. — Heteronucia venusta Nobili, 1906a: 260; 1907: 379, pl. 5, fig. 14 ("Ohura" = Hao). — FOREST &
GUuINOT, 1961: 13, fig. 3a-b, 4, pl. 5, fig. 1-2 (Fakahina; Syn.); 1962: 56 (Biogeography "Tahiti-Tuamotu"). — PEYROT-
CLAUSADE, 1977a, annex of the species: 25; 1977b: 212 (Moorea); 1989: 113 (Tikehau). — GUINOT, 1985: 453 (List).
— Nucia gelida Rathbun, 1907: 68, pl. 5, fig. 4, pl. 9, fig. 2 (Fakarava). — GUINOT, 1985: 453 (List).
Nucia rosea Nobili, 1906
DISTRIBUTION. — Gambier (Mangareva).
REFERENCES. — Nucia rosea Nobili, 1906a: 261; 1907: 381 (Mangareva). — FOREST & GUINOT, 1962: 56
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List).
Nursia mimetica Nobili, 1906
DISTRIBUTION. — Gambier (Mangareva).
REFERENCES. — Nursia mimetica Nobili, 1906a: 261; 1907: 380, pl. 5, fig. 13 (Mangareva). — FOREST & GUINOT,
1962: 56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List).
FAMILY MAJIDAE
Acanthophrys cristimanus A. Milne Edwards, 1865
DISTRIBUTION. — Marquesas (Nuku Hiva).
REFERENCES. — Acanthophrys cristimanus A. Milne Edwards, 1865a: 141, pl. 5, fig. 3 (Nuku Hiva). — GRIFFIN &
TRANTER, 1986: 105 (cf. Remark).
REMARK. — GRIFFIN & TRANTER (1986) have re-established the validity of the genus Acanthophrys, formerly
transferred in Hyastenus, and Acanthophrys cristimanus is the type species of the genus.
Camposcia retusa Latreille, 1829
DISTRIBUTION. — Tuamotu (Fangataufa) - Littoral to deep?
REFERENCES. — Camposcia retusa - GUINOT, 1985: 452, with a ? (List) - NEW MATERIAL - Coll. and det. J. POUPIN
(Fangataufa; 220m, cf. Remark).
REMARK. — The presence of this species in French Polynesia, only inferred by GUINOT because of its large distribution,
is here confirmed by one specimen, collected at an unusual depth.
Cyclax suborbicularis (Stimpson, 1858)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Cyclax suborbicularis - FOREST & GUINOT, 1961: 15, fig. 5-6, 8 bis, 10, pl. 6, fig. 1-2 (Tahiti); 1962:
56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List).
27
Huenia proteus de Haan, 1839
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Huenia proteus - KROpP & BIRKELAND, 1981: 630, tab. 5 (Moorea). — GUINOT, 1985: 452 (List).
Hyastenus aff. borradailei (Rathbun, 1907)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Hyastenus aff. borradaeilli (sic) - PEYROT-CLAUSADE, 1989: 115 (Moorea).
Menaethius monoceros (Latreille, 1825)
DISTRIBUTION. — Marquesas; Society (Moorea, Tahiti); Tuamotu (Fakarava, Hao, Raroia, Takapoto, Tikehau) - Littoral
to sublittoral (30m).
REFERENCES. — Menaethius tuberculatus Dana, 1852b: 123; 1855, pl. 5, fig. 1a-c (Tuamotu). — Menaethius
monoceros - NOBILI, 1907: 382 ("Ohura" = Hao). — RATHBUN, 1907: 64 (Fakarava). — HOLTHUIS, 1953: 4 (Raroia).
— ForEST & GUINOT, 1961: 14, fig. 9a-b (Tahiti; Syn.); 1962: 56 (Biogeography "Tahiti-Tuamotu, Marquesas"). —
PEYROT-CLAUSADE, 1977a, annex of the species: 25 (Moorea); 1977b: 212; 1989: 112, 115 (Moorea, Tikehau; 30m). —
ODINETZ, 1983: 208 (Moorea, Tahiti, Takapoto). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE
FORGES, 1985: 201 (Moorea and/or Tahiti, Takapoto).
Micippa margaritifera Henderson, 1893
DISTRIBUTION. — Tuamotu (Tikehau).
REFERENCES. — Miccipa (sic) margaritifera - PEYROT-CLAUSADE, 1989: 112 (Tikehau).
Micippa parca Alcock, 1895
DISTRIBUTION. — Tuamotu (Makemo).
REFERENCES. — Lophomicippa limbata Rathbun, 1907: 65, pl. 5, fig. 3, pl. 6, fig. 1, 1g (Makemo). — FOREST &
GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List). — Micippa parca - GRIFFIN &
TRANTER, 1986: 277 (Syn.).
Micippoides angustifrons A. Milne Edwards, 1873
DISTRIBUTION. — Tuamotu (Raroia).
REFERENCES. — Micippoides angustifrons - HOLTHUIS, 1953: 5 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST
& GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List).
Perinea tumida Dana, 1851
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava, Takapoto, Tikehau).
REFERENCES. — Perinea tumida - RATHBUN, 1907: 65 (Fakarava). — FOREST & GUINOT, 1962: 56 (Biogeography
"Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 25; 1977b: 212 (Moorea); 1989: 112, 115
(Moorea, Tikehau). — KROPP & BIRKELAND, 1981: 630, tab. 5 (Moorea, Takapoto). — ODINETZ, 1983: 208 (Moorea,
Tahiti, Takapoto). — GUINOT, 1985: 453 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or
Tahiti, Takapoto).
Schizophrys aspera (H. Milne Edwards, 1834)
DISTRIBUTION. — French Polynesia.
REFERENCES. — Schizophrys aspera - GUINOT, 1985: 453, with a ? (List).
REMARK. — Cited by GUINOT, only because of the large distribution of the species (see SAKAI, 1976, or DAI & YANG,
1991: Japon, Hawaii, ... Australia).
28
Simocarcinus obtusirostris (Miers, 1879)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Trigonothir obtusirostris - FOREST & GUINOT, 1961: 14 (Tahiti); 1962: 56 (Biogeography "Tahiti-
Tuamotu"). — Simocarcinus obtusirostris - GUINOT, 1985: 453 (List). — GRIFFIN & TRANTER, 1986: 98 (Syn.).
Tylocarcinus dumerilii (H. Milne Edwards, 1834)
DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Hao).
REFERENCES. — Tylocarcinus gracilis - NOBILI, 1907: 382 (Hao, Mangareva). — FOREST & GUINOT, 1962: 56
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). — Tylocarcinus dumerilii - GRIFFIN & TRANTER,
1986: 197, fig. 67a-b (Syn.) - SYNONYMS - Tylocarcinus gracilis Miers, 1879.
FAMILY PARTHENOPIDAE
Actaeomorpha alvae Boone, 1934
DISTRIBUTION. — Society (Raiatea).
REFERENCES. — Acfaeomorpha alvae Boone, 1934: 37 pl. 11 (Raiatea). — FOREST & GUINOT, 1962: 56 (Biogeography
"Tahiti-Tuamotu”). — GUINOT, 1966b: 759 (cf. Remark); 1985: 453 (List).
REMARK. — According to the drawing published by BOONE (1934), GUINOT (1966b) considers that this species could
possibly be, either Actaemorpha erosa Miers, 1878, or A. punctata Edmonson, 1935.
We keep here the genus Actaeomorpha in the Parthenopidae, but GUINOT (1966b, 1967), in her study of the genera
Aethra (cf. hereafter A. scruposa), Osachila, Hepatus, Hepatella and Actaeomorpha, has modified this usual
classification and placed these genera in a group /ncertae sedis Parthenoxystomata (cf. GUINOT's, 1985 List).
Aethra scruposa (Linné, 1764)
DISTRIBUTION. — Society.
REFERENCES. — Aethra scruposa - GUINOT, 1985: 453, with a ? (Society; listed only according to the large distribution
of the species, without material from French Polynesia).
Daldorfia horrida (Linné, 1758)
DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti); Tuamotu (Fangatau, Hao, Mataiva).
REFERENCES. — Parthenope horrida - NOBILI, 1907: 382 ("Fagatau" = Fangatau, Hao, Mangareva). — SEURAT, 1934:
60 ("Fagatau" = Fangatau, Gambier, Hao). — CHABOUIS L. & F., 1954: 91, fig. 7 (Tahiti, Tuamotu). —FOREST &
GUINOT, 1961: 26, fig. 14 (Tahiti); 1962: 58 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 174, annex 1,
tab. a (Mataiva). — GUINOT, 1985: 453 (List). — Daldorfia horrida - SAKAI, 1976: 283, pl. 96, fig. 2, text-fig. 157
(Syn.).
Parthenope contrarius (Herbst, 1796)
DISTRIBUTION. — Marquesas (Eiao) - Sublittoral.
REFERENCES. — Parthenope contrarius - NEW MATERIAL - Coll. and det. J. POUPIN (Eiao; 42m).
REMARK. — This new material (1 ? and 3 juveniles, Marara st. D38) agrees very well with the description and the
good photograph published by RATBHUN (1906: 885, pl. 17, fig. 1), under Parthenope (Rhinolambrus) lamelligera
(White, 1847). According to SAKAI (1976: 273) WHITE's species is a synonym of P. (Rhinolambrus) pelagicus
(Riippell, 1830), but RATHBUN's material belongs to P. contrarius.
Parthenope hoplonotus (Adams & White, 1848)
DISTRIBUTION. — Society (Tahiti).
Wg)
REFERENCES. — Aulacolambrus hoplonotus - FOREST & GUINOT, 1961: 26, fig. 12a-c, 13 (Tahiti); 1962: 58
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). — Parthenope (Aulacolambrus) hoplonotus - SAKAI,
1976: 280 (Syn.)
FAMILY EUMEDONIDAE
Echinoecus pentagonus (A. Milne Edwards, 1879)
DISTRIBUTION. — Tuamotu (Hao, Raroia).
REFERENCES. — Eumedon convictor Bouvier & Seurat, 1905: 629 (Hao). — NoBILI, 1907: 382 (Hao). — SEURAT,
1934: 58 (Hao). — Eumedonus convictor - HOLTHUIS, 1953: 6 (Raroia). — MORRISON, 1954: 6 (Raroia) —
Echinoecus pentagonus - SERENE et al., 1958: 152 (Syn.). — GUINOT, 1985: 453 (List). — Eumedonus pentagonus -
FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu").
FAMILY PORTUNIDAE
SUBFAMILY CATOPTRINAE
Carupa tenuipes Dana, 1852
DISTRIBUTION. — Gambier (Akamaru); Society (Huahine, Maiao?, Moorea, Tahiti); Tuamotu (Makatea, Makemo,
Marutea South, Pukapuka, Raroia, Tikehau).
REFERENCES. — Carupa tenuipes Dana, 1852a: 85 (Tuamotu); 1852b: 279; 1855, pl.17, fig. 4a-e ("Paumotu
archipelago?”). — A. MILNE EDWARDS, 1861: 386 (Tuamotu; DANA's material). — STEPHENSON & REES, 1967: 5
(Huahine, Moorea, Maiai = Maiao?, "Tickahau" = Tikehau). — SAKAI, 1976: 325 (Syn.). — STEPHENSON, 1976: 12
(Pukapuka). — MONTEFORTE, 1984: 173, annex 1, tab. a (Makatea, Moorea); 1987: 8 (Moorea). — GUINOT, 1985: 449
(List). — Carupa laeviuscula Heller, 1862: 520 (Tarti" = Tahiti); 1865: 27, pl.3, fig. 2 (Tahiti). — NoBILI, 1907: 386
(Akamaru, "Marutea-Vaitutaki" = Marutea South). — RATHBUN, 1907: 64 (Makemo). — HOLTHUIS, 1953: 9 (Raroia).
— Morrison, 1954: 13 (Raroia). — FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu") - NEW MATERIAL -
coll. C. HiLy, det. K. MoosA (Tikehau).
Catoptrus nitidus A. Milne Edwards, 1870
DISTRIBUTION. — Marquesas (Eiao); Society (Huahine); Tuamotu (Makemo, Tikehau) - Sublittoral.
REFERENCES. — Catoptrus nitidus - RATHBUN, 1907: 60 (Makemo). — FOREST & GUINOT, 1962: 58 (Biogeography
"Tahiti-Tuamotu"). — STEPHENSON, 1972: 29 (Syn.). — GUINOT, 1985: 449 (List). — Libystes truncatifrons -
STEPHENSON & REES, 1967: 6 ("Maroe" = Huahine, "Tickahau" = Tikehau) - NEW MATERIAL - Coll. J. POUPIN, det. K.
Moosa (Eiao; 42m) - SYNONYMS - Libystes truncatifrons (de Man, 1887).
SUBFAMILY CAPHYRINAE
Caphyra rotundifrons (A. Milne Edwards, 1869)
DISTRIBUTION. — Society (Bora Bora, Tahiti).
REFERENCES. — Caphyra rotundifrons - RATHBUN, 1907: 60, pl. 1, fig. 4 (Tahiti). — ForREsT & GUINOT, 1962: 58
(Biogeography "Tahiti-Tuamotu”). — STEPHENSON & REES, 1967: 7 (Bora Bora, "Mata Uta Papeete" = Tahiti). —
GUINOT, 1985: 449 (List).
30
Caphyra tridens Richters, 1880
DISTRIBUTION. — Gambier (Mangareva).
REFERENCES. — Caphyra rotundifrons var. tridens Richters - NOBILI, 1907: 386 (""Rikitea, Teone Kura" = Mangareva).
— Caphyra tridens - CROSNIER, 1975: 747, fig. 3a-n (Mangareva).
Lissocarcinus elegans Boone, 1934
DISTRIBUTION. — Society (Raiatea).
REFERENCES. — Lissocarcinus elegans Boone, 1934: 50, pl. 16 (Raiatea). — FOREST & GUINOT, 1962: 58
(Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 27 (Distribution; French Polynesia only). — GUINOT, 1985:
449 (List).
Lissocarcinus laevis Miers, 1886
DISTRIBUTION. — Marquesas (Eiao, Hiva Oa, Nuku Hiva) - Sublittoral.
REFERENCES. — Lissocarcinus laevis - STEPHENSON, 1976: 12 (Nuku Hiva) - NEW MATERIAL - Coll. J. POUPIN, det. K.
Moosa (Eiao, Hiva Oa; 42-53m).
Lissocarcinus orbicularis Dana, 1852
DISTRIBUTION. — Society (Huahine, Moorea, Raiatea, Tahiti); Tuamotu (Hikueru, Moruroa, Tikehau).
REFERENCES. — Lissocarcinus orbicularis - FOREST & GUINOT, 1961: 27, fig. 15a-b, 16a-c (Hikueru); 1962: 58
(Biogeography "Tahiti-Tuamotu"). — STEPHENSON & REES, 1967: 7 (Huahine, Raiatea, "Tikahau" = Tikehau). —
STEPHENSON, 1976: 12 (Moorea). — CHEVALIER et al., 1968: 112, 137 (Moruroa). — MONTEFORTE, 1984: 173, annex
1, tab. a (Tahiti). — GUINOT, 1985: 449 (List). — PEYROT-CLAUSADE, 1989: 113 (Tikehau).
SUBFAMILY PORTUNINAE
Charybdis annulata (Fabricius, 1798)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Goniosoma annulatum (Fabricius) - ORTMANN, 1893a: 82 (Tahiti). — Charybdis annulata - FOREST
& GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). — Charybdis (Charybdis) annulata - CROSNIER, 1962: 78, fig.
136-139, pl. 5, fig. 2 (Distribution only, Tahiti). — GUINOT, 1985: 449, with a ? (List).
Charybdis erythrodactyla (Lamarck, 1818)
DISTRIBUTION. — Austral (Rurutu); Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Tahiti); Tuamotu
(Fangataufa, Hikueru, Makatea, Mataiva, Moruroa, Raroia, Taiaro, Takapoto).
REFERENCES. — Goniosoma erythrodactylum - A. MILNE EDWARDS, 1861: 369 (Marquesas). — DE MAN, 1889: 424
(Tahiti). — ORTMANN, 1893a: 81 (Marquesas). — Charybdis erythrodactyla - NOBILI, 1906b: 118, fig. 3 (Nuku Hiva).
— SENDLER, 1923: 40 (Makatea). — BOONE, 1934: 57, pl. 18-19 (Tahiti). — LEENE, 1936: 117, fig. 1-5 (Marquesas,
Makatea). — HOLTHUIS, 1953: 6 (Raroia). — MORRISON, 1954: 16 (Raroia). — FOREST & GUINOT, 1961: 30 (Hikueru);
1962: 58 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1966a: 48 (Raroia). — CHEVALIER et al., 1968: 92,
137 (Fangataufa). — MONTEFORTE, 1984: 173, annex 1, tab. a, photograph p. 140(b) (Makatea, Mataiva, Takapoto). —
DELESALLE, 1985: 289 (Mataiva). — SALVAT, 1986b: 72, photograph (French Polynesia). — Charybdis
(Goniosupradens) erythrodactyla - LEENE, 1938: 134, fig. 77-80 (Marquesas). — STEPHENSON & REES, 1967: 13 (Bora
Bora, Moorea). — STEPHENSON, 1976: 15 (""Taiohae" = Nuku Hiva). — GUINOT, 1985: 449 (List). — POUPIN 1994a:
29, fig. 25, pl. 3d (Rurutu, Taiaro) - NEW MATERIAL - Coll. and det. J. POUPIN (Moruroa).
Charybdis hawaiensis Edmonson, 1954
DISTRIBUTION. — Tuamotu (Taiaro).
eee NNER aN NTE lala viens Mr
Sil
REFERENCES. — Charybdis (Charybdis) hawaiensis - POUPIN, 1994a: 28, fig. 24, pl. 3c (Taiaro).
Charybdis orientalis Dana, 1852
DISTRIBUTION. — Society.
REFERENCES. — Charybdis (Charybdis) orientalis - LEENE, 1938: 69 (Society; cf. Remark).
REMARK. — Although LEENE (1938) does not mention clearly the location "Society", she indicates that F. CHACE has
examined, for her, in the USNM collections, a male from the Society Islands attributed to C. orientalis (confronted with
the type specimen).
Charybdis paucidentata A. Milne Edwards, 1861
DISTRIBUTION. — Marquesas (Hiva Oa, Tahuata); Tuamotu (Taiaro) - Littoral to sublittoral.
REFERENCES. — Charybdis (Gonioinfradens) paucidentata - POUPIN, 1994a: 30, fig. 26, pl. 3e (Hiva Oa, Tahuata,
Taiaro; 0-100m); 1996: in press (Hiva Oa, Tahuata).
Lupocyclus quinquedentatus Rathbun, 1906
DISTRIBUTION. — Austral (Maria, Rurutu); Marquesas (Nuku Hiva); Society (Bora Bora) - Sublittoral to deep.
REFERENCES. — Lupocyclus quinquedentatus - STEPHENSON, 1976: 15 ("“Hatwata" = Haatuatua bay, Nuku Hiva) - NEw
MATERIAL - Coll. J. POUPIN, det. K. MoosA (Bora Bora, Maria, Rurutu; 80-110m).
Portunus alexandri (Rathbun, 1907)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Callinectes alexandri Rathbun, 1907: 61, pl. 2, fig. 1, pl. 9, fig. 3, 3a-b (Tahiti). — GUINOT, 1985: 449
(List).
REMARK. — This portunid has been described from Tahiti, with a paratype from "Suva, Fidjis". According to
STEPHENSON (1976: 13), it is in fact a non-identifiable Portunus.
Portunus dubius (Laurie, 1906)
DISTRIBUTION. — Marquesas (Eiao, Fatu Hiva, Hiva Oa, Nuku Hiva, Tahuata) - Sublittoral to deep.
REFERENCES. — Portunus dubius - STEPHENSON, 1976: 16 (Marquesas; numerous stations without precisions) - NEW
MATERIAL - Coll. J. POUPIN, det. K. MOoSA (Eiao, Fatu Hiva, Hiva Oa, Nuku Hiva, Tahuata; 42-140m).
Portunus granulatus (H. Milne-Edwards, 1834)
DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Tahiti); Tuamotu
(Fakarava, Raroia, Tikehau).
REFERENCES. — Neptunus (Achelous) granulatus - ORTMANN, 1893a: 72 (Tahiti). — NOBILI, 1907: 383 ("Rikitea" =
Mangareva). — BOONE, 1934: 60, pl. 20 (Nuku Hiva). — SEURAT, 1934: 59 (Mangareva). — Portunus (Achelous)
granulatus - RATHBUN, 1907: 60 (Bora Bora, Fakarava). — SENDLER, 1923: 40 (Tahiti). — Portunus (Cycloachelous)
granulatus - HOLTHUIS, 1953: 6 (Raroia). — MORRISON, 1954: 7 (Raroia). — GUINOT, 1985: 449 (List). — Portunus
granulatus - FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu, Marquesas"). — STEPHENSON & REES,
1967: 25 (Moorea, Tuamotu). — TURKAY, 1971: 127 (Tahiti). — STEPHENSON, 1976: 16 (Tahiti). — MONTEFORTE,
1984: 173, annex 1, tab. a (Moorea, Tahiti); 1987: 8 (Moorea) - NEW MATERIAL - Coll. C. HILY, det. K. Moosa (Tahiti,
Tikehau).
Portunus guinotae Stephenson & Rees, 1961
DISTRIBUTION. — Tuamotu (Marutea South).
REFERENCES. — Portunus guinotae Stephenson & Rees, 1961: 425, fig. 1b, d, g, 2d-f (Marutea South). — FOREST &
GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 39 (Marutea South). — Portunus
(Xiphonectes) guinotae - GUINOT, 1985: 449 (List).
32
Portunus iranjae Crosnier, 1962
DISTRIBUTION. — Marquesas (Eiao, Fatu Hiva, Hiva Oa, Tahuata); Society (Moorea) - Littoral to sublittoral.
REFERENCES. — Portunus iranjae - STEPHENSON & REES, 1967: 30 (""Papetoai bay" = Moorea). — STEPHENSON, 1976:
16 ("Marquesas expedition, st. THX" = Haava strait, between Tahuata and Hiva Oa, cf. HARALD, 1967). — Portunus
(Xiphonectes) iranjae - GUINOT, 1985: 449 (List) - NEW MATERIAL - Coll. J. POUPIN, det. K. MOOSA (Eiao, Fatu Hiva,
Hiva Oa, Tahuata; 54m).
Portunus longispinosus (Dana, 1852)
DISTRIBUTION. — Marquesas (Hiva Oa, Tahuata); Tuamotu (Marutea South, Raroia).
REFERENCES. — Neptunus (Hellenus) longispinosus - NoBILI, 1907: 383 (Marutea South). — Portunus (Hellenus)
longispinosus - HOLTHUIS, 1953: 7 (Raroia). — MORRISON, 1954: 8 (Raroia). — Portunus longispinosus - FOREST &
GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1976: 16 ("Marquesas expedition st. THX, haul
5" = Haava strait, between Tahuata and Hiva Oa, cf. HARALD, 1967). — Portunus (Xiphonectes) longispinosus -
GUINOT, 1985: 449 (List).
REMARK. — This species can be confused with all the species belonging to the Jongispinosus complex, as Portunus
iranjae and P. macrophthalmus (cf. STEPHENSON & REES, 1967; STEPHENSON, 1976; and NAGAI, 1981).
Portunus macrophthalmus Rathbun, 1906
DISTRIBUTION. — Marquesas (Eiao, Hiva Oa) - Sublittoral.
REFERENCES. — Portunus macrophthalmus - NEW MATERIAL - Coll. J. POUPIN, det. K. Moos (Eiao, Hiva Oa; 42-
53m).
Portunus nipponensis Sakai, 1938
DISTRIBUTION. — Tuamotu (Moruroa) - Sublittoral to deep.
REFERENCES. — Portunus nipponensis - POUPIN et al. 1990: 17 (French Polynesia). — POUPIN, 1996: in press pro
parte (Moruroa; cf. Remark).
REMARK. — This species is usually found in shallow waters (15-50m) but, in French Polynesia, it has been trapped up
to 130m. Except for Moruroa, the localities mentioned in POUPIN (1996) concerned in fact a new species, related to P.
nipponensis, but with a distinct male pleopod (Moosa & CROSNIER, in study).
Portunus orbitosinus Rathbun, 1911
DISTRIBUTION. — Marquesas (Eiao) - Sublittoral.
REFERENCES. — Portunus orbitosinus - NEW MATERIAL - Coll. J. POUPIN, det. K. MoosA (Eiao; 42m).
Portunus pelagicus (Linné, 1758)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Neptunus pelagicus - HELLER, 1865: 27 (Tahiti). — Portunus pelagicus - FOREST & GUINOT, 1962:
58 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 41 (Distribution only, Tahiti). — Portunus (Portunus)
pelagicus - GUINOT, 1985: 449 (List).
REMARK. — Although this species has often been reported from Tahiti (BOONE, 1934; STEPHENSON & CAMPBELL, 1959;
CROSNIER, 1962; STEPHENSON & REES, 1967; STEPHENSON, 1972; SAKAI, 1976; DAI & YANG, 1991), it seems that the
single material examined from this locality is mentioned in HELLER (1865). It would thus be very interesting to check
this reference.
Portunus sanguinolentus (Herbst, 1783)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Mataiva).
REFERENCES. — Neptunus sanguinolentus - CHABOUIS L. & F., 1954: 91, fig. 9 (French Polynesia). — Portunus
sanguinolentus - FOREST & GUINOT, 1961: 29, fig. 17a-b, 18 (Tahiti); 1962: 58 (Biogeography "Tahiti-Tuamotu"). —
a RN OSES RPL TORO SU SSE ee
33
MONTEFORTE, 1984: 173, annex 1, tab. a (Moorea). — SALVAT, 1986b: 72 (French Polynesia). — POUPIN, 1994a: 31,
fig. 27, pl. 3f (Mataiva, Tahiti). — Portunus sanguinolentus sanguinolentus - STEPHENSON, 1976: 19 (Tahiti). —
Portunus (Portunus) sanguinolentus - GUINOT, 1985: 449 (List).
Scylla serrata (Forskal, 1775)
DISTRIBUTION. — Society (Huahine, Raiatea, Tahiti, Tupai) - Brackish to sea water.
REFERENCES. — Scylla serrata - HELLER, 1865: 27 (Tahiti). — MIERS, 1886: 185 (Tahiti). — BOONE, 1934: 68, pl. 25-
30 (Huahine, Tahiti). — SEURAT, 1934: 58 (Tahiti). — CHABOUIS L. & F., 1954: 90, unnumbered fig. (Huahine,
Raiatea). — FOREST & GUINOT, 1961: 27 (Tahiti); 1962: 58 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48
(Society); 1985: 449 (List). — BABLET, 1972: 32, pl. 11 (French Polynesia). — SALVAT, 1986b: 70, 72 (French
Polynesia) - NEW MATERIAL - Coll. J. POUPIN, det. K. MoosA (Raiatea, Tupai).
Thalamita admete (Herbst, 1803)
DISTRIBUTION. — Gambier (Mangareva); Marquesas?; Society (Bora Bora, Huahine, Moorea, Raiatea, Tahiti);
Tuamotu (Fakarava, Hikueru, Kaukura, Makemo, Mataiva, Takapoto, Tikehau).
REFERENCES. — Thalamita admete - HELLER, 1865: 28 (Tahiti). — ORTMANN, 1893a: 83 (Tahiti). — NOBILI, 1907:
383 (Kaukura, Mangareva). — RATHBUN, 1907: 63 (Fakarava, Makemo). — FOREST & GUINOT, 1961: 30, fig. 19a-b
(Hikueru, Tahiti); 1962: 58 (Biogeography "Tahiti-Tuamotu, Marquesas" with a ?). — STEPHENSON & REES, 1967: 56,
fig. 20 (Bora Bora, Huahine, Moorea, Raiatea, Tahiti, "Tikahau" = Tikehau). — NAIM, 1980a, annex 1, tab. 3 (Moorea).
— MONTEFORTE, 1984: 173, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 8 (Moorea). — DELESALLE,
1985: 288 (Mataiva). — Thalamita (Thalamita) admete - GUINOT, 1985: 449 (List).
Thalamita bouvieri Nobili, 1906
DISTRIBUTION. — Gambier (Mangareva).
REFERENCES. — Thalamita bouvieri Nobili, 1906a: 262; 1907: 384, pl. 2, fig. 2 ("Rikitea" = Mangareva). — SEURAT,
1934: 60 (French Polynesia). — CROSNIER, 1962: 119, fig. 201-204, pl. 10, fig. 2 (Mangareva; syntypes). — FOREST &
GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — Thalamita (Pseudothalamitopsis) bouvieri - GUINOT, 1985:
449 (List; subgenus from Moosa, 1979: 47) - RELEVANT MATERIAL - Thalamitoides (sic) aff. bouvieri - PEYROT-
CLAUSADE, 1989: 113 (Tikehau).
Thalamita chaptalti (Audouin, 1826)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Thalamita chaptalii - FOREST & GUINOT, 1961: 34, fig. 21a-b (Tahiti); 1962: 60 (Biogeography
"Tahiti-Tuamotu"). — Thalamita (Neothalamita) chaptalii - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979:
43).
Thalamita coerulipes Jacquinot, 1852
DISTRIBUTION. — Gambier (Kamaka, Mangareva); Society (Bora Bora, Huahine, Tahiti); Tuamotu (Fakarava).
REFERENCES. — Thalamita coerulipes Jacquinot, 1852, pl. 5, fig. 6-10 (Mangareva). — JACQUINOT & LUCAs, 1853: 53
(Mangareva). — A. MILNE EDWARDS, 1861: 363 (Mangareva). — NOBILL, 1907: 383 (Kamaka) — RATHBUN, 1907: 63
(Fakarava, Society). — BOONE, 1934: 78, pl. 35 (Bora Bora). — FOREST & GUINOT, 1961: 32 (Tahiti); 1962: 60
(Biogeography "Tahiti-Tuamotu"). — STEPHENSON & REES, 1967: 64 (Huahine). — POUPIN, 1994a: 32, fig. 28, pl. 3g
(Kamaka, Mangareva, Tahiti, Tuamotu). — Thalamita (Thalaminella) coerulipes - GUINOT, 1985: 449 (List; subgenus
from Moosa, 1979: 51).
Thalamita cooperi Borradaile, 1903
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Thalamita cooperi - NAIM, 1980a: annex 1, tab. 3 (Moorea; material not found in MNHN).
34
Thalamita corrugata Stephenson & Rees, 1961
DISTRIBUTION. — Tuamotu (Tikehau).
REFERENCES. — Thalamita corrugata - STEPHENSON & REES, 1967: 65, fig. 23 ("Tickahau" = Tikehau). — GUINOT,
1985: 449 (List).
Thalamita crenata (Latreille, 1829)
DISTRIBUTION. — Marquesas; Society (Bora Bora, Maiao?, Moorea, Tahiti); Tuamotu (Mataiva, Takapoto, Tikehau).
REFERENCES. — Thalamita crenata - A. MILNE EDWARDS, 1861: 365 (Marquesas). — ORTMANN, 1893a: 86
(Marquesas). — RATHBUN, 1907: 62 (Bora Bora). — SEURAT, 1934: 59 (Marquesas). — FOREST & GUINOT, 1962: 60
(Biogeography "Tahiti-Tuamotu, Marquesas"). — STEPHENSON & REES, 1967: 66 ("Maiai" = Maiao?, "Tikahau" =
Tikehau). — MONTEFORTE, 1984: 173, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 8 (Moorea). —
SALVAT & RICHARD, 1985: 356, 362 (Takapoto) — Thalamita (Thalaminella) crenata - GUINOT, 1985: 449 (List;
subgenus from Moosa, 1979: 51) - NEW MATERIAL - Coll. C. HILY, det. K. Moosa (Tahiti, Tikehau).
Thalamita dakini Montgomery, 1931
DISTRIBUTION. — Society (Bora Bora, Moorea); Tuamotu (Makatea, Mataiva, Takapoto).
REFERENCES. — Thalamita dakini - STEPHENSON & REES, 1967: 69 (Bora Bora, Moorea). — MONTEFORTE, 1984: 173,
annex 1, tab a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — Thalamita (Thalamitopsis)
dakini - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 47).
Thalamita danae Stimpson, 1858
DISTRIBUTION. — French Polynesia.
REFERENCES. — Thalamita danae - NEW MATERIAL - Coll. PLEssIs, det. K. MoosA (French Polynesia).
Thalamita demani Nobili, 1905
DISTRIBUTION. — Marquesas (Nuku Hiva).
REFERENCES. — Thalamita demani - STEPHENSON, 1976: 20 (""Hatwata" = Haatuatua bay, Nuku Hiva).
Thalamita edwardsi Borradaile, 1900
DISTRIBUTION. — Gambier (Mangareva, Temoe); Society (Tahiti).
REFERENCES. — Thalamita admete var. Edwardsii Bort. - NOBILI, 1907: 383 ("Rikitea" = Mangareva, "Timoe" =
Temoe). — Thalamita edwardsi - FOREST & GUINOT, 1961: 32, fig. 20a-b (Tahiti); 1962: 58 (Biogeography "Tahiti-
Tuamotu"). — Thalamita edwardsi - GUINOT, 1985: 449, with " =T. admete ?" (List; cf. Remark).
REMARK. — This species was formerly considered as a synonym of Thalamita admete by STEPHENSON & HUDSON
(1957). This assertion is not followed by FOREST & GUINOT (1961), CROSNIER (1962) and DAI & YANG (1991).
Thalamita gatavakensis Nobili, 1906
DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Tahiti).
REFERENCES. — Thalamita pilumnoides var. gatavakensis Nobili, 1906a: 262; 1907: 384 (""Gatavake" = Mangareva).
— Thalamita pilumnoides ssp. gatavakensis - FOREST & GUINOT, 1961: 34, fig. 22a-b, 23-25 (Mangareva); 1962: 60
(Biogeography "Tahiti-Tuamotu"). — Thalamita gatavakensis - CROSNIER, 1962: 99, fig. 156a-c, e (Mangareva;
syntypes). — STEPHENSON & REES, 1967: 75 (Bora Bora, Tahiti). — Thalamita (Thalamita) gatavakensis - GUINOT,
1985: 449 (List) - NEW MATERIAL - Coll. C. HILY, det. K. Moosa (Tahiti).
Thalamita gloriensis Crosnier, 1962
DISTRIBUTION. — Society (Huahine).
35
REFERENCES. — Thalamita gloriensis - STEPHENSON & REES, 1967: 76 (Huahine). — Thalamita (Thalamita)
gloriensis - GUINOT, 1985: 449 (List).
Thalamita gracilipes (A. Milne Edwards, 1873)
DISTRIBUTION. — Society (Tahiti); Tuamotu (Moruroa?).
REFERENCES. — Thalamita gracilipes - STEPHENSON, 1976: 21 (Tahiti) - RELEVANT MATERIAL - Thalamonyx aff.
gracilipes - SALVAT & RENAUD-MORNANT, 1969: 165 (Moruroa). — Thalamita (Thalamonyx) aff. gracilipes -
GUINOT, 1985: 449 (List).
Thalamita integra Dana, 1852
DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti); Tuamotu (Takapoto, Nukutipipi).
REFERENCES. — Thalamita integra Dana, 1852a: 85; 1852b: 281; 1855, pl. 17, fig. 6a-d (Tuamotu). — A. MILNE
EDWARDS, 1861: 358 (Tahiti, Tuamotu). — NoBIL, 1907: 383 ("Gatavake" = Mangareva). — FOREST & GUINOT, 1962:
60 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 173, annex 1, tab. a (Takapoto). — SALVAT & RICHARD,
1985: 350 (Takapoto). — MERSCHARDT-SALVAT, 1991: 89 (Nukutipipi). — Thalamita (Thalamita) integra - GUINOT,
1985: 449 (List).
Thalamita macropus Montgomery, 1931
DISTRIBUTION. — Austral (Neilson bank); Marquesas (Fatu Hiva) - Sublittoral to deep.
REFERENCES. — Thalamita macropus - NEW MATERIAL - Coll. B. RICHER DE FORGES and J. POUPIN, det. K. MOOSA
(Neilson bank, Fatu Hiva; 49-100/130m).
Thalamita macrospinifera Rathbun, 1911
DISTRIBUTION. — Austral (Rurutu); Marquesas (Eiao, Hiva Oa); Society (Raiatea); Tuamotu (Makemo, Moruroa) -
Sublittoral to deep.
REFERENCES. — Thalamita macrospinifera - POUPIN, 1996: in press (Makemo, Raiatea; 120-160m) - NEW MATERIAL - F
Coll. J. POUPIN, det. K. Moosa (Eiao, Hiva Oa, Makemo, Moruroa, Raiatea, Rurutu; 80-160m).
Thalamita minuscula Nobili, 1906
DISTRIBUTION. — Tuamotu (Kaukura, Vahitahi).
REFERENCES. — Thalamita minuscula Nobili, 1906a: 262; 1907: 386, pl. 1, fig. 15 (Kaukura, Vahitahi). — FOREST &
GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 49 (List with "Only from Tuamotu Is.").
— Thalamita (Neothalamita) minuscula - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 43).
REMARK. — This very small species (maximum width, 4mm), never recorded since its description, could be the juvenile
of another species.
Thalamita mitsiensis Crosnier, 1962
DISTRIBUTION. — Austral (Rurutu) - Sublittoral.
REFERENCES. — Thalamita mitsiensis - NEW MATERIAL - Coll. J. POUPIN, det. K. MOOSA (Rurutu; 80m).
Thalamita philippinensis Stephenson & Rees, 1967
DISTRIBUTION. — Austral (Rurutu); Tuamotu (Moruroa) - Sublittoral to deep.
REFERENCES. — Thalamita philippinensis - NEW MATERIAL - Coll. J. POUPIN, det. K. Moosa (Moruroa, Rurutu; 95-
130m).
Thalamita picta Stimpson, 1858
DISTRIBUTION. — Marquesas (Hiva Oa, Nuku Hiva, Tahuata); Gambier?; Society (Raiatea, Tahiti); Tuamotu (Fakarava,
Makatea, Mataiva, Raroia, Takapoto).
oe ee ss
36
REFERENCES. — Goniosoma lineatum A. Milne Edwards, 1861: 377, pl. 35, fig. 4 (Nuku Hiva) fide CROSNIER (1962:
138). — Thalamita alcocki - NOBILI, 1907: 384 (""Tagatau" = Gambier?). — Thalamita gardineri - RATHBUN, 1907: 63
(Fakarava, Makemo). — Thalamita picta - HOLTHUIS, 1953: 8 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST &
GUINOT, 1961: 33 (Tahiti; Syn.); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 50 (Syn.); 1976:
23 ("Hoava Strait” = Haava strait, between Tahuata and Hiva Oa, cf. HARALD, 1967). — MONTEFORTE, 1984: 173,
annex 1, tab. a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — Thalamita (Neothalaminella)
picta - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 51) - NEW MATERIAL - Coll. J. POUPIN, det. K. MOOSA
(Raiatea) - SYNONYMS - Thalamita alcocki de Man, 1902; T. gardineri Borradaile, 1902.
Thalamita pilumnoides Borradaile, 1903
DISTRIBUTION. — Society (Huahine, Moorea, Raiatea).
REFERENCES. — Thalamita pilumnoides - STEPHENSON & REES, 1967: 87, fig. 32 (Huahine, Moorea, Raiatea). —
PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212 (Moorea). — MONTEFORTE, 1984: 173, annex 1, tab.
a; 1987: 8 (Moorea). — Thalamita (Neothalamita) pilumnoides - GUINOT, 1985: 449 (List; subgenus from Moosa,
1979: 43).
Thalamita prymna (Herbst, 1803)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Anaa).
REFERENCES. — Thalamita prymna - STEPHENSON, 1976: 23 (Anaa). — PEYROT-CLAUSADE, 1989: 115 (Moorea) -
NEW MATERIAL - Coll. J. POUPIN, det. K. MOoSA (Tahiti).
Thalamita quadrilobata Miers, 1884
DISTRIBUTION. — Society (Bora Bora).
REFERENCES. — Thalamita quadrilobata - STEPHENSON & REES, 1967: 92 (Bora Bora). — Thalamita
(Pseudothalamitopsis) quadrilobata - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 47).
Thalamita seurati Nobili, 1906
DISTRIBUTION. — Tuamotu (Marutea South).
REFERENCES. — Thalamita seurati Nobili, 1906a: 262; 1907: 385, pl. 2, fig. 1 (Marutea). — SEURAT, 1934: 60 (French
Polynesia). — FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 51 (List
"Tuamotu"). — Thalamita (Pseudothalamitopsis) seurati - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979:
47).
Thalamita spinifera Borradaile, 1903
DISTRIBUTION. — Austral (Raevavae, Tubuai); Marquesas (Eiao, Fatu Hiva, Nuku Hiva); Tuamotu (Makemo) -
Sublittoral to deep.
REFERENCES. — Thalamita spinifera - STEPHENSON, 1976: 24 ("Marquesas Expedition, 40-80m, 18/ix/1967 to
1/x/1967" = Marquesas, cf. HARALD, 1967) - NEW MATERIAL - Coll. J. POUPIN, det. K. MOOSA (Eiao, Fatu Hiva,
Makemo, Nuku Hiva, Raevavae, Tubuai; 42-200m).
Thalamita woodmasoni Alcock, 1899
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Thalamita woodmasoni - FOREST & GUINOT, 1961: 33 (Tahiti); 1962: 60 (Biogeography "Tahiti-
Tuamotu"). — Thalamita (Pseudothalamitopsis) woodmasoni - GUINOT, 1985: 449 (List; subgenus from Moosa,
1979: 47).
Thalamitoides quadridens A. Milne Edwards, 1869
DISTRIBUTION. — Tuamotu (Moruroa, Rangiroa).
EE EEE eee
37
REFERENCES. — Thalamitoides quadridens - STEPHENSON, 1976: 26 (Rangiroa) - NEW MATERIAL - Coll. J. POUPIN, det.
K. Moosa (Moruroa).
SUBFAMILY PODOPHTHALMINAE
Podophthalmus vigil (Fabricius, 1798)
DISTRIBUTION. — Society (Moorea, Tahiti).
REFERENCES. — Podophthalmus vigil - FOREST & GUINOT, 1961: 36 (Tahiti); 1962: 60 (Biogeography "Tahiti
Tuamotu"). — STEPHENSON & REES, 1967: 104 ("Opunohu Bay” = Moorea). — MONTEFORTE, 1984: 173, annex 1, tab.
a (Moorea). — GUINOT, 1985: 449 (List).
FAMILY XANTHIDAE
SUBFAMILY POLYDECTINAE.
Lybia caestifera (Alcock, 1897)
DISTRIBUTION. — Society (Tahiti); Tuamotu (Mataiva?).
REFERENCES. — Lybia caestifera - RATHBUN, 1907: 60 (Tahiti). — FoREST & GUINOT, 1962: 68 (Biogeography
"Tahiti-Tuamotu"). — GUINOT, 1976: 75 (Syn.; RATHBUN's reference with a ?); 1985: 452, with a ? (List) - RELEVANT
MATERIAL - Lybia cf. coestifera (sic) - MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva).
Lybia plumosa Barnard, 1947
DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau).
REFERENCES. — Lybia leptochelis - PEYROT-CLAUSADE, 1977a, annex of the species: 27 (Moorea) not L. leptochelis
(Zehntner, 1894) = L. plumosa fide SERENE (1984: 29, 31). — Lybia plumulosa (sic) - PEYROT-CLAUSADE, 1989: 113
(Tikehau).
Lybia tessellata (Latreille, 1812)
DISTRIBUTION. — Marquesas; Society (Moorea, Tahiti); Tuamotu (Makatea, Mataiva, Rangiroa, Raroia).
REFERENCES. — Melia tesselata - FINNEGAN, 1931: 647 (Marquesas). — Lybia tessellata - HOLTHUIS, 1953: 23
(Raroia). — Morrison, 1954: 13 (Raroia). — FOREST & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu,
Marquesas”). — TAKEDA & MIYAKE, 1970: 15 ("Avatoru” = Rangiroa, Tahiti). — GUINOT, 1976: 70, fig. 17d, 18e, 19c,
20e-h, 22d, pl. 2, fig. 6 (Syn.); 1985: 452 (List). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva,
Moorea, Tahiti); 1987: 9 (Moorea).
Polydectus cupulifer (Latreille, 1812)
DISTRIBUTION. — Tuamotu (Raraka).
REFERENCES. — Polydectus villosus Dana, 1852a: 81; 1852b: 227; 1855, pl. 13, fig. 3a-e (Raraka). — Polydectus
cupulifer - FOREST & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1976: 65 (Syn.); 1985: 452
(List). — SERENE, 1984: 24, fig. 1, pl. 1a (Syn.).
38
SUBFAMILY CYMOINAE
Cymo andreossyi (Audouin, 1826)
DISTRIBUTION. — Society (Moorea, Tahiti).
REFERENCES. — Cymo andreossyi - DANA, 1852b: 225; 1855, pl. 13, fig. 2a-b (Tahiti). — HELLER, 1865: 20 (Tahiti).
— Forest & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu"), — ODINETZ, 1983: 206 (Moorea, Tahiti). —
GUINOT, 1985: 450 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti).
Cymo deplanatus A. Milne Edwards, 1873
DISTRIBUTION. — Tuamotu (Raroia).
REFERENCES. — Cymo deplanatus - HOLTHUIS, 1953: 18 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST &
GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 450 (List).
Cymo melanodactylus de Haan, 1833
DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Hao, Hikueru,
Moruroa).
REFERENCES. — Cymo Andreossyi var. melanodactyla - NOBILI, 1907: 397 ("Otepa" = Hao, Mangareva). — Cymo
melanodactylus - RATHBUN, 1907: 53 (Bora Bora, Fakarava). — PESTA, 1913: 46 (Tahiti). — BOONE, 1934: 144, pl. 74
(Tahiti). — FOREST & GUINOT, 1961: 119 (Hikueru, "Rikitea" = Mangareva); 1962: 68 (Biogeography "Tahiti-
Tuamotu"). — ODINETZ, 1983: 206 (Tahiti). — SERENE, 1984: 34, fig. 8, pl. 2b (Syn.). — GUINOT, 1985: 450 (List). —
ODINETZ-COLLART & RICHER DE ForRGES, 1985: 201 (Tahiti). — POUPIN, 1994a: 33, fig. 29, pl. 3h (Mangareva,
Moruroa).
Cymo quadrilobatus Miers, 1884
DISTRIBUTION. — Society (Moorea, Tahiti?).
REFERENCES. — Cymo quadrilobatus - ODINETZ, 1983: 206 (Moorea). — GUINOT, 1985: 450 (List). — ODINETZ-
COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti).
SUBFAMILY TRICHIINAE
Banareia parvula (Krauss, 1843)
DISTRIBUTION. — Marquesas.
REFERENCES. — Actaea parvula - ODHNER, 1925: 35, 51, pl. 3, fig. 13 (Marquesas). — Banareia parvula - GUINOT,
1976: 179, with a ? for the genus (Syn.; see the considerations about the generic rank); 1985: 452 (List). — Banareia
parvula - GARTH et al., 1987: 243 (cited only for the generic rank).
SUBFAMILY LIOMERINAE
Liomera bella (Dana, 1852)
DISTRIBUTION. — Gambier (Mangareva, Tarauru-Roa, Vaiatekeue); Society (Moorea, Tahiti); Tuamotu (Fakareva,
Hikueru, Makatea, Makemo, Marutea South, Mataiva, Takapoto, Takaroa, Tikehau).
REFERENCES. — Carpiloxanthus rugipes - HELLER, 1865: 17 (Tahiti) not Liomera rugipes (Heller, 1861) = L. bella fide
SERENE (1984: 65). — Carpilodes rugatus - NoBILI, 1907: 387 ("chenal Waiatekene = Vaiatekeue, "Rikitea” =
Mangareva, "Waitutaki" = Marutea South). — RATHBUN, 1907: 37 (Makemo) - All, not Liomera rugata (H. Milne
Edwards, 1834) = L. bella fide SERENE (1984: 61). — Carpilodes vaillantianus - NOBILI, 1907: 387. — Carpilodes
bellus - ODHNER, 1925: 16, pl. 1, fig. 9 ("Eimeo" = Moorea, Makemo, Tahiti). — BUITENDIJK, 1960: 257, fig. 2b
(Tahiti). — Liomera bella - FOREST & GUINOT, 1961: 38, fig. 26a-b (Hikueru, Tahiti, "Taraourou-roa" = Tarauru-Roa);
eT
39
1962: 60 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212
(Moorea); 1989: 111 (Tikehau). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti,
Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 450 (List). — ODINETZ-COLLART
& RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti). — Liomera (Liomera) bella - SERENE, 1984: 60, fig. 21, pl.
5e (Syn.) - SYNONYMS - Carpilodes vaillantianus A. Milne Edwards, 1862.
Liomera cinctimana (White, 1847)
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Moorea, Tahiti).
REFERENCES. — Liomera lata - HELLER, 1865: 9 (Tahiti). — Liomera cinctimana - ORTMANN, 1893b: 450, pl. 17, fig.
8 (Tahiti). — FOREST & GUINOT, 1961: 39, fig. 27a-b (Tahiti, "Taihoae” = Nuku Hiva); 1962: 60 (Biogeography
"Tahiti-Tuamotu, Marquesas"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26 (Moorea). — MONTEFORTE,
1984: 171, annex 1, tab. a (Tahiti). — GUINOT, 1985: 450 (List). — Carpilodes cinctimanus - ODHNER, 1925: 14
(Tahiti). — Liomera (Liomera) cinctimana - SERENE, 1984: 57, fig. 17, pl. Sa (Syn.) - SYNONYMS - Liomera lata Dana,
1852.
Liomera laevis (A. Milne Edwards, 1873)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Liomera laevis - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212 (Moorea). —
GUINOT, 1985: 451, with a ? (List).
Liomera laperousei Garth, 1985
DISTRIBUTION. — Austral (MacDonald bank) - Littoral to sublittoral .
REFERENCES. — Liomera laperousei - LABOUTE & RICHER DE FORGES, 1986: 21 (MacDonald bank, 40m; with
hesitation: "semble étre une femelle juvénile de Liomera laperousei Garth, 1985 décrit de lille de Paque").
Liomera monticulosa (A. Milne Edwards, 1873)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Marutea South) - Littoral to sublittoral.
REFERENCES. — Carpilodes monticulosus - NOBILI, 1907: 387 (Marutea South). — ODHNER, 1925: 21, pl. 1, fig. 18
(Tahiti, Marutea South). — Liomera monticulosa - FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). —
GuINoT, 1964: 11 (Syn.); 1985: 451 (List). — PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212
(Moorea); 1989: 115 (Moorea; 30m). — Liomera (Liomera) monticulosa - SERENE, 1984: 64, fig. 24, pl. 6c (Syn.). —
Not Carpilodes monticulosus - RATHBUN, 1907: 37 (Fakarava, Makemo) = Liomera (Liomera) rugata (H. Milne
Edwards, 1834) fide SERENE (1984: 62).
Liomera pallida (Borradaile, 1900)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Carpilodes pallidus - ODHNER, 1925: 20, pl. 1, fig. 17 (Tahiti). — Liomera pallida - FOREST &
GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 451, with a ? (List). — Liomera (Liomera)
pallida - SERENE, 1984: 62, pl. 5f (Syn.).
Liomera rubra (A. Milne Edwards, 1865)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Liomera rubra - MONTEFORTE, 1984: 171, annex 1, tab. a; 1987: 9 (Moorea). — GUINOT, 1985: 451,
with a ? (List). — Liomera (Liomera) rubra - SERENE, 1984: 65, fig. 26, pl. 6e-f, pl. 9f (Syn.).
Liomera rugata (H. Milne Edwards, 1834)
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Moorea, Tahiti); Tuamotu (Fakarava, Makemo, Mataiva, Raroia,
Tikehau).
40
REFERENCES. — Carpilodes rugatus - ORTMANN, 1893b: 468 (Tahiti). — ODHNER, 1925: 20, pl. 1, fig. 16 (Fakarava,
Tahiti). — BOONE, 1934: 91, pl. 46 (Nuku Hiva). — HOLTHUIS, 1953: 13 (Raroia). — MORRISON, 1954: 16 (Raroia). —
BUITENDUK, 1960: 259, fig. 2d (Tahiti). — Carpilodes monticulosus - RATHBUN, 1907: 37 (Fakarava, Makemo) not
Liomera monticulosa (A. Milne Edwards, 1873) = L. rugata fide SERENE (1984: 62). — Liomera rugata - FOREST &
GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva, Moorea);
1987: 9 (Moorea). — GUINOT, 1985: 451 (List). — PEYROT-CLAUSADE, 1989: 113, 115 (Moorea, Tikehau). — Liomera
(Liomera) rugata - SERENE, 1984: 62, fig. 22, pl. 6b (Syn.).
Liomera semigranosa De Man, 1888
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Liomera semigranulosa (sic) - ODINETZ, 1983: 209 (Tahiti). — Liomera semigranosa - GUINOT,
1985: 451 (List). — Liomera (Liomera) semigranosa - SERENE, 1984: 63, pl. 7c, f (Syn.).
Liomera stimpsoni (A. Milne Edwards, 1865)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Liomera stimpsoni - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212; 1989: 115
(Moorea). — GUINOT, 1985: 451, with a ? (List).
Liomera tristis (Dana, 1852)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava).
REFERENCES. — Carpilodes tristis Dana, 1852a: 77; 1852b: 193; 1855, pl. 9, fig. 7a-d (Tuamotu with a ?). — HELLER,
1865: 17 (Tahiti). — A. MILNE EDWARDS, 1865b: 225 (Tuamotu). — DE MAN, 1890: 50 (Tahiti). — RATHBUN, 1907:
37 (Fakarava). — ODHNER, 1925: 12, pl. 1, fig. 1 ("Eimeo" = Moorea, Tahiti). — BUITENDIJK, 1960: 254, fig. lc
(Tahiti). — Carpilodes granulatus Heller, 1862: 520 (Tahiti). — Liomera tristis - FOREST & GUINOT, 1961: 38
(Tahiti); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 451 (List). — Liomera (Liomera) tristis -
SERENE, 1984: 59, fig. 19, pl. 5b (Syn.).
Liomera venosa (H. Milne Edwards, 1834)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Carpilodes venosus - ORTMANN, 1893b: 467 (Tahiti). — ODHNER, 1925: 22, pl. 2, fig. 1 (Tahiti). —
BUITENDIJK, 1960: 259, fig. 2e (Tahiti). — Liomera venosa - FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti-
Tuamotu"). — GUINOT, 1985: 451 (List). — Liomera (Liomera) venosa - SERENE, 1984: 58, fig. 18, pl. 7d-e (Syn.).
Neoliomera demani Forest & Guinot, 1961
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru).
REFERENCES. — Neoliomera demani Forest & Guinot, 1961: 80, fig. 76, 77bis, pl. 3, fig. 3-5 (Hikueru, Tahiti); 1962:
66 (Biogeography "Tahiti-Tuamotu"). — SERENE, 1984: 71, fig. 31, pl. 8b (Hikueru). — GuINoT, 1985: 451 (List). —
PEYROT-CLAUSADE, 1989: 115 (Moorea). — Neoliomera pubescens - ODHNER, 1925: 28, pl. 2, fig. 6, 6a-7 (Tahiti) not
Neoliomera pubescens (H. Milne Edwards, 1834) = N. demani sp. nov. in FOREST & GUINOT (1961: 80).
Neoliomera insularis (White, 1847)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Neoliomera insularis - SAKAI, 1976: 398 (Tahiti; cf. Remark). — GUINOT, 1985: 451 (List; after
SAKAI).
REMARK. — The only reference in French Polynesia seems to be in SAKAI (1976) where "Tahiti" is cited without
material examined from that island, and without older references for that location. Thus, the occurence of this species in
the French Polynesia still remains to be confirmed.
41
Neoliomera pubescens (H. Milne Edwards, 1834)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Neoliomera pubescens - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212 (Moorea).
— SERENE, 1984: 71, fig. 30, pl. 8a (Syn.; cf. Remark).
REMARK. — According to SERENE (1984) it is, in most of the works, necessary to check that the material attributed to
Neoliomera pubescens has not been confounded with N. demani Forest & Guinot, 1961. He also mentions that N.
pubescens is known, with certainty, only from Maunitius.
Neoliomera richtersi (De Man, 1889)
DISTRIBUTION. — Society (Tahiti); Tuamotu (Marutea South).
REFERENCES. — Actaeodes richtersii de Man 1889: 412, pl. 9, fig. 2; 1890: 51 (Tahiti). — Liomera richtersi - NOBILI,
1907: 387 (""Marutea-Vaitutaki" = Marutea South). — Neoliomera richtersii - ODHNER, 1925: 33, pl. 2, fig. 13 (Tahiti).
— BUITENDIJK, 1960: 262 (Tahiti). — FOREST & GUINOT, 1961: 79, fig. 74 (Tahiti); 1962: 66 (Biogeography "Tahiti-
Tuamotu"). — GUINOT, 1964: 47, fig. 17 (Tahiti); 1985: 451 (List). — SERENE, 1984: 70, fig. 28, pl. 8e (Tahiti; Syn.).
Neoliomera variolosa (A. Milne Edwards, 1873)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Neoliomera variolosa - PEYROT-CLAUSADE, 1977b: 212, 220 (Moorea).
SUBFAMILY EUXANTHINAE
Alainodaeus rimatara Davie, 1993
DISTRIBUTION. — Austral (Raevavae, Rimatara); Tuamotu (Akiaki, Fangataufa, Hao, Takapoto) - Sublittoral to deep.
REFERENCES. — Alainodaeus rimatara Davie, 1993: 519, fig. 6, pl. 6 (Akiaki, Fangataufa, Hao, Raevavae, Rimatara,
Takapoto; 90-350m). — POUPIN, 1996: in press (same material).
Euxanthus exsculptus (Herbst, 1790)
DISTRIBUTION. — Gambier (Mangareva); Society (Moorea, Tahiti).
REFERENCES. — Euxanthus exsculptus var. rugosus - NOBILI, 1907: 389 (Mangareva) not Euxanthus rugosus Miers,
1884 = E. exsculptus fide GUINOT-DUMORTIER (1960b: 170). — Euxanthus exsculptus - GUINOT-DUMORTIER, 1960b:
169, pl. 1, fig. 4, pl. 2, fig. 10, pl. 6, fig. 36-37, pl. 8, fig. 42-47 (""Rikitea" = Mangareva). — FOREST & GUINOT, 1962:
62 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, annex 1, tab. a; 1987: 9 (Moorea). — SERENE, 1984:
86, fig. 48, pl. 11b (Tahiti). — GuINOT, 1985: 450 (List).
Euxanthus sculptilis Dana, 1852
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Euxanthus sculptilis - BOONE, 1934: 107, pl. 57 (Tahiti). — ForREsT & GUINOT, 1962: 62
(Biogeography "Tahiti-Tuamotu"). — GUINOT-DUMORTIER, 1960b: 167, pl. 6, fig. 39, pl. 9, fig. 49 (Syn.); 1985: 450
(List).
Medaeus grandis Davie, 1993
DISTRIBUTION. — Tuamotu (Hao, Moruroa) - Sublittoral to deep.
REFERENCES. — Medaeus grandis Davie, 1993: 526, fig. 8, pl. 8 (Hao, Moruroa; 90-210m). — PouPIN, 1996: in press
(same material).
Paramedaeus noelensis (Ward, 1934)
DISTRIBUTION. — Society (Moorea, Tahiti).
42
REFERENCES. — Medaeus noelensis - FOREST & GUINOT, 1961: 56, fig. 42-43, 44a-b, pl. 1, fig. 1 (Tahiti); 1962: 62
(Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a (Moorea, Tahiti); 1987: 9 (Moorea). —
Paramedaeus noelensis - SERENE, 1984: 90, fig. 51, pl. 12f (Syn.). — GUINOT, 1985: 451 (List).
Paramedaeus simplex (A. Milne Edwards, 1873)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Paramedaeus simplex - PEYROT-CLAUSADE, 1977b: 212 (Moorea). — GUINOT, 1985: 451, with a ?
(List).
SUBFAMILY ACTAEINAE
Actaea aff. glandifera Rathbun, 1914
DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral.
REFERENCES. — Actaea aff. glandifera - PEYROT-CLAUSADE, 1989: 111, 114 (Moorea, Tikehau; 25m).
Actaea calculosa (H. Milne Edwards, 1834)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Actaea calculosa - ODHNER, 1925: 52 (Tahiti). — FOREST & GUINOT, 1962: 64 (Biogeography "Tahiti-
Tuamotu").
REMARK. — ODHNER mentions a Tahitian specimen in the collections of Hamburg. However, GUINOT (1976: 215-216)
states that most of the references to Actaea calculosa, especially ODHNER (1925: 52), must be re-examined.
Actaea danae A. Milne Edwards, 1865
DISTRIBUTION. — Tuamotu (Raraka).
REFERENCES. — Actaeodes areolatus Dana, 1852a: 77; 1852b: 194; 1855, pl. 9, fig. 8a-d (Raraka). — Actaea danae -
ForEST & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — (Actaea) danae - GUINOT, 1976: 247 (Syn.); 1985:
450, with "species inquirenda" (List).
Actaea polyacantha (Heller, 1861)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Actaea polyacantha - PEYROT-CLAUSADE, 1989: 114 (Moorea).
Actaeodes consobrinus (A. Milne Edwards, 1873)
DISTRIBUTION. — Marquesas; Society (Moorea) - Littoral to sublittoral.
REFERENCES. — Actaea consobrina - ODHNER, 1925: 67, pl. 4, fig. 14 (Marquesas). — Actaeodes consobrinus -
GUINOT, 1976: 246, pl. 15, fig. 5, 5a (Syn.); 1985: 450, with a ? (List). — Actaeodes consobrina - PEYROT-CLAUSADE,
1989: 114 (Moorea; 30m). — Not Actaea consobrina - NOBILI, 1907: 390 = Actaea ruppellioides sp. nov. in ODHNER
(1925: 47; cf. under Pseudoliomera ruppellioides).
Actaeodes hirsutissimus (Riippell, 1830)
DISTRIBUTION. — Society (Bora Bora, Moorea, Raiatea, Tahiti); Tuamotu (Mataiva, Tikehau).
REFERENCES. — Actaea hirsutissima - HELLER, 1865: 9 (Tahiti). — RATHBUN, 1907: 42 (Bora Bora, Tahiti). —
ODHNER, 1925: 69, pl. 4, fig. 13 (Tahiti). — BOONE, 1934: 124, pl. 66 (Raiatea). — FOREST & GUINOT, 1961: 78
(Tahiti); 1962: 64 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1989: 111 (Tikehau). — Actaeodes
hirsutissimus - GUINOT, 1976: 245, fig. 38e, pl. 15, fig. 2, 2a (Tahiti; Syn.); 1985: 450 (List). — PEYROT-CLAUSADE,
1977a, annex of the species: 26; 1977b: 213; 1985: 462 (Moorea). — NAIM, 1980a, annex 1, tab. 3 (Moorea). —
MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva, Moorea, Tahiti); 1987: 8 (Moorea). — SERENE, 1984: 135 (Syn.).
eee
43
Actaeodes tomentosus (H. Milne Edwards, 1834)
DISTRIBUTION. — Society (Tahiti); Tuamotu (Makatea).
REFERENCES. — Actaeodes tomentosus - HELLER, 1865: 17 (Tahiti). — GUINOT, 1976: 244, fig. 38d, 41c, pl. 15, fig. 1,
la (Syn.); 1985: 450, with a ? (List). — SERENE, 1984: 134, 137 (Syn.). — Actaea tomentosa - SENDLER, 1923: 37
(Makatea). — FoREST & GUINOT, 1962: 66 (Biogeography "Tahiti-Tuamotu”).
Forestia depressa (White, 1847)
DISTRIBUTION. — Marquesas.
REFERENCES. — Actaea depressa - BALSS, 1935: 136 (Marquesas). — Forestia depressa - GUINOT, 1976: 262 (Syn.,;
with a ? for BALSS' reference); 1985: 450 (List; with a ? in front of "Marquesas"). — SERENE, 1984: 106 (Syn.; with a ?
in front of BALSs' reference).
REMARK. — In GUINOT (1976) and SERENE (1984), at least one specimen attributed to Actaea depressa, by BALSS
(1938: 54), would be in fact a Forestia scabra (Odhner, 1925).
Forestia scabra (Odhner, 1925)
DISTRIBUTION. — Marquesas.
REFERENCES. — Actaea scabra - BALSS, 1935: 136 (Marquesas). — Forestia scabra - GUINOT, 1976: 263 (Syn.; with a
? in front of BALSS' reference); 1985: 450 (List; with a ? in front of "Marquesas").
Gaillardiellus rueppelli (Kraus, 1843)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Gaillardiellus rueppelli - GARTH & KIM, 1983: 684 (Distribution only, Tahiti; cf. Remark). —
GUINOT, 1985: 450 (List; presumably after the previous work).
REMARK. — The material examined by GARTH & KIM (1983: 685) was collected in the Philippines by the Albatross
(1908-1909). "Tahiti", mentioned in the “Distribution” only, corresponds neither to material examined, nor to former
references cited in this work.
Gaillardiellus superciliaris (Odhner, 1925)
DISTRIBUTION. — Tuamotu (Raroia, Taiaro).
REFERENCES. — Actaea superciliaris - HOLTHUIS, 1953: 11 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST &
GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — Gaillardiellus superciliaris - GUINOT, 1976: 257 (Syn.); 1985:
450 (List). — POUPIN, 1994a: 34, fig. 30, pl. 4a, with a ? (Taiaro).
Paractaea excentrica Guinot, 1969
DISTRIBUTION. — Tuamotu (Marutea South).
REFERENCES. — Paractaea excentrica Guinot, 1969: 263, fig. 36 (Marutea South); 1985: 451 (List).
Paractaea retusa (Nobili, 1905)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Raroia).
REFERENCES. — Actaea garretti Rathbun, 1906: 852, pl. 9, fig. 8 (Society). — Actaea rufopunctata - HOLTHUIS, 1953:
11 (Raroia). — Morrison, 1954: 16 (Raroia). — FOREST & GUINOT, 1961: 79, fig. 79a-b (Tahiti) - All, not Paractaea
rufopunctata (H. Milne Edwards, 1834) = Paractaea retusa form hippocrepica nov. in GUINOT (1969: 256). —
Paractaea retusa form hippocrepica - GUINOT, 1969: 256, fig. 30 (Tahiti, Raroia; Syn.); 1985: 451 (List). — PEYROT-
CLAUSADE, 1989: 115 (Moorea). — Paractaea retusa - MONTEFORTE, 1984: 171, annex 1, tab. a; 1987: 9 (Moorea).
REMARK. — GUINOT (1969: 255), when creating the new genus Paractaea, has examined the syntype of garretti from
Gilbert Islands (but not the specimen from the Society Islands), and considers that RATHBUN's species belongs to
Paractaea retusa (Nobili) form garretti (Rathbun).
Paractaea rufopunctata H. Milne Edwards, 1834
DISTRIBUTION. — Society (Tahiti); Tuamotu (Makemo, Marutea South).
REFERENCES. — Actaea rufopunctata - NOBILI, 1907: 392 (Marutea). — RATHBUN, 1907: 43 (Makemo, Tahiti). —
ODHNER, 1925: 60 (Tahiti). — FOREST & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — Paractaea
rufopunctata form plumosa - GUINOT, 1969: 248, fig. 21 (Marutea). — Paractaea rufopunctata - GUINOT, 1985: 451
(List).
Paractaeopsis quadriareolatus (Takeda & Miyake, 1968)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Paractaeopsis quadriareolatus - SERENE, 1984: 127 (Syn.; gen. nov.). — Paractaea quadriareolata -
PEYROT-CLAUSADE, 1989: 115 (Moorea).
Paractaeopsis tumulosus (Odhner, 1925)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Actaea tumulosa Odhner, 1925: 61, pl. 4, fig. 10 (Tahiti). — ForEST & GUINOT, 1962: 66
(Biogeography "Tahiti-Tuamotu”). — Paractaeopsis tumulosus - SERENE, 1984: 127, fig. 74, pl. 17d (Syn.; gen. nov.).
— Paractaea tumulosa - GUINOT, 1985: 451 (List).
Psaumis cavipes (Dana, 1852)
DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Hao, Rangiroa,
Raroia, Tikehau).
REFERENCES. — Actaea cavipes - NOBILI, 1907: 390 ("Ohura" = Hao, "Rikitea" = Mangareva). — RATHBUN, 1907: 44,
pl. 1, fig. 2 (Bora Bora, Fakarava, Rangiroa). — ODHNER, 1925: 68 ("Eimeo" = Moorea, Tahiti). — BOONE, 1934: 128,
pl. 68 (Tahiti). — HOLTHUIS, 1953: 10 (Raroia). — MORRISON, 1954: 16 (Raroia). — FOREST & GUINOT, 1961: 78
(Tahiti); 1962: 64 (Biogeography "Tahiti-Tuamotu”). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b:
212 (Moorea). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a (Moorea, Tahiti);
1987: 8 (Moorea). — Psaumis cavipes - ODINETZ, 1983: 209 (Moorea, Tahiti). — SERENE, 1984: 129, fig. 76, pl. 18f
(Syn.). — GUINOT, 1985: 451 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti,
Takapoto). — MONTEFORTE, 1987: 9 (Moorea). — PEYROT-CLAUSADE, 1989: 113 (Tikehau).
Psaumis cellulosa (Dana, 1852)
DISTRIBUTION. — Society (Moorea, Tahiti).
REFERENCES. — Psaumis cellulosa - ODINETZ, 1983: 209 (Moorea, Tahiti). — GUINOT, 1985: 451 (List).
Pseudoliomera granosimana (A. Milne Edwards, 1865)
DISTRIBUTION. — Society (Tahiti); Tuamotu (Marutea South).
REFERENCES. — Liomera granosimana - ORTMANN, 1893b: 451 (Tahiti). — NOBILI, 1907: 387 ("Marutea-Vaitutaki" =
Marutea South). — Pseudoliomera granosimana - ODHNER, 1925: 79, fig. 5-6 (Tahiti). — FOREST & GUINOT, 1961:
39, fig. 28a-c, pl. 7, fig. 1-2 (Tahiti); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — SERENE, 1984: 100, fig. 56, pl. 13a
(Tahiti). — GuINOT, 1985: 451 (List).
Pseudoliomera lata (Borradaile, 1902)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Pseudoliomera lata - KROpP & BIRKELAND, 1981: 630, tab. 5 (Moorea). — SERENE, 1984: 102 (Syn.).
— PEYROT-CLAUSADE, 1989: 115 (Moorea). — (Pseudoliomera) lata - GUINOT, 1985: 451, with a ? (List). — Not
Actaea lata - NOBILI, 1907: 392 ("Marutea, Vaitutaki" = Marutea South) = A. ruppellioides Odhner, 1925 fide GUINOT
(1962: 237).
45
Pseudoliomera ruppellioides (Odhner, 1925)
DISTRIBUTION. — Tuamotu (Marutea South).
REFERENCES. — Actaea consobrina - NOBILI, 1907: 390 (Marutea South) not Actaea consobrina A. Milne Edwards,
1873 =A. ruppellioides sp. nov. in ODHNER (1925: 47). — Actaea lata - NoBILI, 1907: 392 ("Marutea, Vaitutaki" =
Marutea South) not Actaea lata Borradaile, 1902 = A. ruppellioides Odhner fide GUINOT (1962: 237). — Actaea
ruppellioides Odhner, 1925: 47, pl. 3, fig. 9 (Marutea South; NOBILI's material). — FOREST & GUINOT, 1962: 64
(Biogeography "Tahiti-Tuamotu"). — (Pseudoliomera) ruppellioides - GUINOT, 1976: 203, 246 (Genus Pseudoliomera
"ou a sa proximité"); 1985: 451 (List).
Pseudoliomera speciosa (Dana, 1852)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Actaea speciosa - SENDLER, 1923: 38 (Tahiti). — ODHNER, 1925: 62 (Tahiti). — FOREST & GUINOT,
1962: 66 (Biogeography "Tahiti-Tuamotu"). — (Pseudoliomera) speciosa - GUINOT, 1976: 203, 243 (Genus uncertain);
1985: 451, with a ? (List).
Pseudoliomera variolosa (Borradaile, 1902)
DISTRIBUTION. — Society (Moorea); Tuamotu (Mataiva, Tikehau) - Littoral to sublittoral.
REFERENCES. — Pseudoliomera variolosa - KROPP & BIRKELAND, 1981: 630, tab. 5 (Moorea). — MONTEFORTE, 1984:
171, annex 1, tab. a (Mataiva, Moorea; cf. Remark); 1987: 9 (Moorea). — PEYROT-CLAUSADE, 1989: 111, 115 (Moorea,
Tikehau; 30m). — Aff. Pseudoliomera variolosa - PEYROT-CLAUSADE, 1977a, annex of the species: 27 (Moorea). —
(Pseudoliomera) variolosa - GUINOT, 1985: 451 (List).
REMARK. — MONTEFORTE writes, Pseudoliomera variolosa (A. Milne Edwards, 1837 sic). Then, it could be
Neoliomera variolosa (A. Milne Edwards, 1873), which is different from BORRADAILE's species (cf. SERENE, 1984: 66).
SUBFAMILY ZOZIMINAE
Atergatis floridus (Linné, 1767)
DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Hao).
REFERENCES. — Atergatis floridus - DANA, 1852b: 159; 1855, pl. 7, fig. 4 (Society and Tuamotu). — HELLER, 1865: 8
(Tahiti). — NosIL, 1907: 388 ("Ohura"” = Hao). — SEURAT, 1934: 59 (Hao). — BUITENDIJK, 1960: 268 (Society). —
FoREST & GUINOT, 1961: 41 (Tahiti; Syn.); 1962: 62 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a,
annex of the species: 26; 1977b: 212 (Moorea). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — MONTEFORTE, 1984: 170,
annex 1, tab. a (Moorea, Tahiti); 1987: 8 (Moorea). — GUINOT, 1985: 450 (List). — PoUPIN, 1994a: 35, fig. 31, pl. 4b
(Mangareva, Tahiti). — Afergatis ocyroe - RATHBUN, 1907: 37 (Bora Bora). — SENDLER, 1923: 37 (Tahiti) -
SYNONYMS - Altergatis ocyroe (Herbst, 1801).
Atergatopsis cf. germaini A. Milne Edwards, 1865
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Atergatopsis cf. germaini - MONTEFORTE, 1984: 170, annex 1, tab. a; 1987: 9 (Moorea). — GUINOT,
1985: 450 (List).
Atergatopsis signatus (Adams & White, 1848)
DISTRIBUTION. — Tuamotu (Makatea, Mataiva, Raroia, Takapoto).
REFERENCES. — Atergatopsis signatus - HOLTHUIS, 1953: 12 (Raroia). — MORRISON, 1954: 16 (Raroia). — FOREST &
GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (Raroia); 1985: 450 (List). —
MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). —
SALVAT, 1986b: 72 (French Polynesia).
46
Lophozozymus cristatus A. Milne Edwards, 1867
DISTRIBUTION. — Austral (Maria); Society.
REFERENCES. — Lophozozymus cristatus - BUITENDUK, 1960: 292, fig. 7a (Society). — FOREST & GUINOT, 1962: 62
(Biogeography "Tahiti-Tuamotu"). — POUPIN, 1994a: 36, fig. 32, pl. 4c (Maria).
Lophozozymus dodone (Herbst, 1801)
DISTRIBUTION. — Society (Moorea, Tahiti).
REFERENCES. — Atfergatis elegans Heller, 1862: 519; 1865: 7, pl. 1, fig. 3 (Tahiti). — Lophozozymus dodone - BALSS,
1938: 39 (Tahiti). — FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 171,
annex 1, tab. a (Moorea, Tahiti); 1987: 9 (Moorea). — SERENE, 1984: 171, pl. 24e (Syn.). — GUINOT, 1985: 451 (List).
— Not Lophozozymus dodone - FOREST & GUINOT, 1961: 54, fig. 39a-b (Tahiti) = Lophozozymus glaber fide GUINOT
(1979: 65).
Lophozozymus edwardsi Odhner, 1925
DISTRIBUTION. — Gambier (Mangareva); Society (Moorea); Tuamotu (Marutea South).
REFERENCES. — Lophozozymus superbus - NoBILI, 1907: 388 (Mangareva, "Marutea Vaitutaki" = Marutea South) not
Lophozozymus superbus A. Milne Edwards, 1873 = L. edwardsi fide FOREST & GUINOT (1961: 56) & GuINoT (1979:
63). — Lophozozymus edwardsi - FOREST & GUINOT, 1961: 56, fig. 41 (Mangareva); 1962: 62 (Biogeography "Tahiti-
Tuamotu”). — GUINOT, 1979: 63 (Mangareva, Marutea South); 1985 (List). — MONTEFORTE, 1984: 171, annex 1, tab.
a; 1987: 9 (Moorea).
Lophozozymus glaber Ortmann, 1893
DISTRIBUTION. — Society (Moorea, Tahiti).
REFERENCES. — Lophozozymus glaber - GUINOT, 1979: 65, pl. 8, fig. 2, 2a (Tahiti). — NAIM, 1980a, annex 1, tab. 3
(Moorea). — MONTEFORTE, 1984: 171, annex 1, tab. a; 1987: 9 (Moorea). — Lophozozymus dodone - FOREST &
GUINOT, 1961: 54, fig. 39a-b (Tahiti) not Lophozozymus dodone (Herbst, 1801) = L. glaber fide GUINOT (1979: 65).
Lophozozymus pictor (Fabricius, 1798)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Lophozozymus octodentatus - BOONE, 1934: 102, pl. 54-55 (Tahiti). — SAKAI, 1976: 407, pl. 146, fig.
3 (Syn., but without BOONE's reference) - SYNONYMS - Lophozozymus octodentatus (H. Milne Edwards, 1834).
REMARK. — BOONE has examined and illustrated a large male and a female from Tahiti. However, it seems that this
material has never been re-examined (cf. BUITENDIJK, 1960, or SAKAI, 1976).
Lophozozymus superbus (Dana, 1852)
DISTRIBUTION. — Tuamotu (Moruroa, Raraka, Raroia).
REFERENCES. — Xantho superbus Dana, 1852a: 74; 1852b: 167; 1855, pl. 8, fig. 5a-b (Raraka). — Lophozozymus
superbus - HOLTHUIS, 1953: 23 (Raroia). — MORRISON, 1954: 13 (Raroia). — GUINOT, 1979: 63 (Moruroa); 1985: 451
(List). — Lophozozymus incisus - FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu") not Lophozozymus
incisus (H. Milne Edwards, 1834) = L. superbus (Dana) (cf. Remark). — Not Lophozozymus superbus - NOBILI, 1907:
388 = L. edwardsi fide FOREST & GUINOT (1961: 56) & GUINOT (1979: 63).
REMARK. — FOREST & GUINOT (1962: 62) have recorded Lophozozymus incisus (H. Milne Edwards, 1834) in French
Polynesia because it has formerly been considered as a synonym of L. superbus (Dana). This opinion has been changed
later on (see for example HOLTHUIS, 1953, or GUINOT, 1979).
Platypodia anaglypta (Heller, 1861)
DISTRIBUTION. — Society (Moorea); Tuamotu (Fakarava, Tikehau).
47
REFERENCES. — Platypodia anaglypta - RATHBUN, 1907: 38 (Fakarava). — FOREST & GUINOT, 1962: 62
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 451 (List). — PEYROT-CLAUSADE, 1989: 113, 115 (Moorea,
Tikehau).
Platypodia granulosa (Riippell, 1830)
DISTRIBUTION. — Society (Tahiti?); Tuamotu (Nukutipipi).
REFERENCES. — Atergatis limbatus - ? HELLER, 1865: 8 (Tahiti). — Platypodia granulosa - ? PESTA, 1913: 41 (Tahiti).
— FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — SERENE, 1984: 159, 162 (Syn.). — GUINOT,
1985: 451 (List). — MERSCHARDT-SALVAT, 1991: 89 (Nukutipipi). — Not Lophactea granulosa - NOBILI, 1907: 388
("Rikitea" = Mangareva, Marutea). — Not Platypodia granulosa - FOREST & GUINOT, 1961: 51 (Mangareva) - These
two references = Platypodia pseudogranulosa sp. nov. in SERENE (1984: 159; cf. Remark) - SYNONYMS - Atergatis
limbatus (H. Milne Edwards, 1834).
REMARK. — SERENE (1984: 159), for the description of Platypodia pseudogranulosa, closely related to P. granulosa,
does not mention HELLER's (1865) and PEsTA's (1913) references, neither under P. granulosa, nor under P.
pseudogranulosa. Thus we place these two references under both species, with uncertainty.
Platypodia pseudogranulosa Seréne, 1984
DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti?); Tuamotu (Marutea South).
REFERENCES. — Afergatis limbatus - ? HELLER, 1865: 8 (Tahiti; cf. Remark under P. granulosa). — Lophactaea
granulosa (Riipp.) - NOBILI, 1907: 388 ("Rikitea” = Mangareva, Marutea). — Platypodia granulosa - ? PESTA, 1913:
41 (Tahiti; cf. Remark under P. granulosa). — FOREST & GUINOT, 1961: 51 (Mangareva) - NOBILI's and FOREST &
GUINOT's references, not Platypodia granulosa (Riippell, 1830) = P. pseudogranulosa sp. nov. in SERENE (1984: 159).
— Platypodia pseudogranulosa Seréne, 1984: 159, pl. 22d ("Rikitea” = Mangareva, Marutea; Syn.).
Platypodia semigranosa (Heller, 1861)
DISTRIBUTION. — Society (Moorea) - Littoral to sublittoral.
REFERENCES. — Platypodia semigranosa - SERENE, 1984: 160, fig. 95, pl. 22b (Syn.; with material collected by
PEYROT-CLAUSADE at Madagascar). — PEYROT-CLAUSADE, 1989: 115 (Moorea; 22m).
Zozimus aeneus (Linné, 1758)
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti); Tuamotu (Fakahina, Hikueru, Makatea, Makemo, Mataiva,
Moruroa, Raroia, Taiaro, Takapoto).
REFERENCES. — Zozymus aeneus - DANA, 1852b: 192; 1855, pl. 10, fig. 3a (Tuamotu). — STIMPSON, 1858a: 32 [30];
1907: 42 (Tahiti). — Noi, 1907: 388 (Fakahina). — SEURAT, 1934: 59 (Fakahina, Tahiti). — MONTEFORTE, 1984:
171, annex 1, tab. a, photograph p. 136b (Makatea, Mataiva, Takapoto). —DELESALLE, 1985: 289 (Mataiva). —
SALVAT & RICHARD, 1985: 360 (Takapoto). — SALVAT, 1986b: 72 (French Polynesia). — Zozimus aeneus - RATHBUN,
1907: 38 (Makemo). — BOoNng, 1934: 99, pl. 50-53 (Nuku Hiva, Tahiti). — HOLTHUIS, 1953: 27 (Raroia). —
Morrison, 1954: 16 (Raroia). — BUITENDIK, 1960: 284, fig. 6a (Society). — FOREST & GUINOT, 1961: 51 (Hikueru,
Tahiti); 1962: 62 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1966a: 48 (Raroia); 1985: 451.(List). —
BONVALLOT et al., 1994: 140-141, photograph (Tuamotu). — PouPIN, 1994a: 37, fig. 33, pl. 4d (Tahiti, Taiaro). — ?
Zozimus sp. - SALVAT, 1986a: 19, photograph (French Polynesia; det. according to the photograph). — ?
Lophozozymus sp. - BAGNIS & CHRISTIAN, 1983: 110, photograph (Tuamotu; det. according to the photograph) - NEW
MATERIAL - Coll. and det. J. POUPIN (Moruroa).
Zozymodes pumilus (Jacquinot, 1852)
DISTRIBUTION. — Gambier?; Society (Tahiti); Tuamotu (Hikueru, Makemo).
REFERENCES. — Xanthodius cristatus - RATHBUN, 1907: 41 (Makemo). — Zozymodes carinipes - NOBILI, 1907: 388
("Tagatau" = Gambier?) not Zozymodes carinipes Heller, 1861 synonym of Z. xanthoides (Krauss, 1843) = Z. pumilus
fide FOREST & GUINOT (1961: 52). — Zozymodes pumilus - FOREST & GUINOT, 1961: 52, fig. 36a-b (Hikueru); 1962:
48
62 (Biogeography "Tahiti-Tuamotu”). — SERENE, 1984: 153, fig. 90, pl. 14e (Tahiti; Syn.). — GumNot, 1985: 452
(List) - SYNONYMS - Leptodius cristatus Borradaile, 1902.
Zozymodes xanthoides (Krauss, 1843)
DISTRIBUTION. — Tuamotu (Takapoto, Tikehau).
REFERENCES. — Zozymodes xanthoides - MONTEFORTE, 1984: 171, annex 1, tab. a (Takapoto). — GUINOT, 1985: 452,
with a ? (List). — Zozymoides xanthoides (sic) - PEYROT-CLAUSADE, 1989: 113 (Tikehau).
SUBFAMILY XANTHINAE
Lachnopodus bidentatus (A. Milne Edwards, 1867)
DISTRIBUTION. — Society (Moorea, Tahiti).
REFERENCES. — Xantho arcuatus Heller, 1865: 11, pl. 2, fig. 1 (Tahiti). — Lachnopodus bidentatus - FOREST &
GUINOT, 1961: 42, fig. 29-30, 32bis, 33a-b, pl. 7, fig. 1-2 (Tahiti; Syn.); 1962: 62 (Biogeography "Tahiti-Tuamotu"). —
PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab.
a; 1987: 9 (Moorea). — GUINOT, 1985: 250 (List).
Lachnopodus ponapensis (Rathbun, 1907)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Xanthias ponapensis Rathbun, 1907: 44, pl. 7, fig. 5, 5a (Tahiti). — Paraxanthias ponapensis -
FOREST & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 451 (List). — Lachnopodus
ponapensis - SERENE, 1984: 203 (Key).
Lachnopodus subacutus (Stimpson, 1858)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Mataiva).
REFERENCES. — Lachnopodus subacutus - PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212 (Moorea).
— MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva, Moorea, Tahiti); 1987: 9 (Moorea). — SERENE, 1984: 204, fig.
122, pl. 29a (Syn.). — GUINOT, 1985: 450, with a ? (List).
Lachnopodus tahitensis De Man, 1889
DISTRIBUTION. — Society (Tahiti); Tuamotu (Makatea, Raroia, Takapoto).
REFERENCES. — Xantho (Lachnopodus) tahitensis de Man, 1889: 418, pl. 9, fig. 4, 4a; 1890: 52 (Tahiti). —
Lachnopodus tahitensis - HOLTHUIS, 1953: 22 (Raroia). — MORRISON, 1954: 16 (Raroia). — FOREST & GUINOT, 1961:
49 (Tahiti); 1962: 62 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (Raroia); 1985: 450 (List). —
MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Takapoto). — SERENE, 1984: 203, fig. 123, pl. 29d (Tahiti).
Leptodius davaoensis Ward, 1941
DISTRIBUTION. — Tuamotu (Hao, Hikueru, Mataiva, Moruroa, Takapoto).
REFERENCES. — Leptodius leptodon Forest & Guinot, 1961: 65, fig. 55-56, 59a-b, pl. 2, fig. 3 (Hikueru) fide TAKEDA
(1980: 318); 1962: 64 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva,
Takapoto). — GUINOT, 1985: 450 (List). — Leptodius exaratus - NOBILI, 1907: 389 (Hao) not Leptodius exaratus (H.
Milne Edwards, 1834) = L. leptodon nov. in FOREST & GUINOT (1961: 65). — Leptodius davaoensis - TAKEDA, 1980:
318 (Syn.). — POUPIN, 1994a: 38, fig. 34, pl. 4e (Hikueru, Moruroa) - RELEVANT MATERIAL - Leptodius cf. davaoensis
- MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva). — GUINOT, 1985: 450 (List).
Leptodius exaratus (H. Milne Edwards, 1834)
DISTRIBUTION. — Marquesas (Nuku Hiva); Society; Tuamotu (Raroia).
neni iii ieee aaa ee
49
REFERENCES. — Leptodius exaratus - BOONE, 1934: 110, pl. 58 (Nuku Hiva). — FoREsT & GUINOT, 1962: 64
(Biogeography "Tahiti-Tuamotu, ?Marquesas"). — SERENE, 1984: 183, fig. 106, pl. 26a (Syn.). — GUINOT, 1985: 450
(List). — Xantho exaratus - HOLTHUIS, 1953: 27 (Raroia). — MORRISON, 1954: 7 (Raroia). — BUITENDUK, 1960: 331,
fig. 9k-m (Society). — Not Leptodius exaratus (H. Milne Edwards) - NoBILI, 1907: 389 = Leptodius leptodon nov. in
ForEST & GUINOT (1961: 65) synonym of L. davaoensis fide TAKEDA (1980: 318).
Leptodius gracilis (Dana, 1852)
DISTRIBUTION. — Gambier (Rikitea); Tuamotu (Hikueru, Moruroa, Rangiroa, Raroia).
REFERENCES. — Leptodius gracilis - NOBILI, 1907: 389 (Rikitea). — FOREST & GUINOT, 1961: 64, fig. 57, 58a-b, pl. 2,
fig. 4 (Hikueru); 1962: 62 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 450 (List). — PoUPIN, 1994a: 39, fig.
35, pl. 4f (Hikueru, Moruroa). — Xantho gracilis - HOLTHUIS, 1953: 27 (Raroia). — MORRISON, 1954: 7 (Raroia). —
BUITENDUK, 1960: 335 (Rangiroa).
Leptodius sanguineus (H. Milne Edwards, 1834)
DISTRIBUTION. — Gambier (Kamaka, Mangareva, Tarauru-Roa); Marquesas (Nuku Hiva); Society (Bora Bora,
Moorea,Tahiti); Tuamotu (Ahe and/or Manihi, Fakarava, Makatea, Makemo, Marutea South, Mataiva, Rangiroa,
Taiaro, Takapoto).
REFERENCES. — Chlorodius sanguineus - DANA, 1852a: 79; 1852b: 207; 1855, pl. 11, fig. 11a-d ("Waterland' = Ahe
and/or Manihi). — Leptodius sanguineus - NoBILI, 1907: 389 (“Rikitea” = Mangareva, Kamaka, Marutea South). —
RATHBUN, 1907: 39 (Bora Bora, Fakarava, Makatea, Makemo, Mangareva, Nuku Hiva, Rangiroa, Tahiti). — SENDLER,
1923: 37 (Makatea, Tahiti). — BOONE, 1934: 116, pl. 60-61 (Nuku Hiva). — FoREST & GUINOT, 1961: 63, fig. 50a-b
("Gatavake" = Mangareva, Tahiti, "Taraourou-roa” = Tarauru-Roa); 1962: 62 (Biogeography "Tahiti-Tuamotu,
Marquesas"). — MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9
(Moorea). — SERENE, 1984: 185 (Syn.). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 450 (List). — POUPIN,
1994a: 40, fig. 36, pl. 4g (Nuku Hiva, Tahiti, Taiaro). — Xantho sanguineus - BUITENDUK, 1960: 323 (Nuku Hiva).
Lioxanthodes alcocki Calman, 1909
DISTRIBUTION. — Society (Moorea); Tuamotu (Makatea, Mataiva, Takapoto, Tikehau).
REFERENCES. — Lioxanthodes alcocki - MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Moorea,
Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 451 (List). — PEYROT-
CLAUSADE, 1989: 113 (Tikehau).
Macromedaeus crassimanus (A. Milne Edwards, 1867)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Xantho crassimanus - BUITENDLK, 1960: 318, fig. 9c-f (Tahiti). — Leptodius crassimanus - FOREST
& GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — Macromedaeus crassimanus - SERENE, 1984: 179, fig. 103,
pl. 25b (Syn.). — GUINOT, 1985: 451 (List).
Macromedaeus distinguendus (de Haan, 1835)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Macromedaeus distinguendus - KIM, 1973: 630 (Distribution only, Tahiti). — GUINOT, 1985: 451
(Society; List).
REMARK. — GUINOT (1985) has listed this species after FOREST & GUINOT (1961). These authors have actually
examined some Xantho distinguendus de Haan, 1835 (p. 57, under Medaeus noelensis Ward, 1934), but they are from
Hong Kong. It could be that GUINOT refers to KIM (1973), who has quoted "Tahiti" in the distribution of this species.
However, this location concerns neither the material examined, nor one of the references cited by KIM under
Macromedaeus distinguendus. The presence of this species in French Polynesia remains thus doubtful.
50
Macromedaeus nudipes (A. Milne Edwards, 1867)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Xantho nudipes - FOREST & GUINOT, 1961: 59, fig. 47a-b. (Tahiti); 1962: 62 (Biogeography "Tahiti-
Tuamotu"). — Macromedaeus nudipes - SERENE, 1984: 178, fig. 101, pl. 25a (Syn.). — GUINOT, 1985: 451 (List).
Neoxanthops cavatus (Rathbun, 1907)
DISTRIBUTION. — Tuamotu (Fakarava).
REFERENCES. — Cycloxanthops cavatus Rathbun, 1907: 41, pl. 5, fig. 8, pl. 6, fig. 3, 3a (Fakarava). — FOREST &
GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — Neoxanthops cavatus - SERENE, 1984: 212, fig. 128, pl. 29f
(Syn.). — GUINOT, 1985: 451 (List).
Paraxanthias notatus (Dana, 1852)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava, Makatea, Makemo, Marutea North?, Marutea South,
Mataiva, Takapoto, Tikehau).
REFERENCES. — Xanthodes notatus Dana, 1852a: 76 (Tahiti, Tuamotu); 1852b: 178; 1855, pl. 8, fig. 12a-b. (Society or
Tuamotu). — Xanthias notatus - NOBILI, 1907: 392 (Fakarava, Makatea, Marutea = Marutea North?, Marutea South).
— RATHBUN, 1907: 45 (Fakarava, Makemo). — Paraxanthias notatus - FOREST & GUINOT, 1961: 76, fig. 70a-b
(Tahiti); 1962: 64 (Biogeography "Tahiti-Tuamotu”). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva,
Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 451 (List). —
PEYROT-CLAUSADE, 1989: 113, 115 (Moorea, Tikehau).
Xanthias canaliculatus Rathbun, 1907
DISTRIBUTION. — Society (Moorea); Tuamotu (Makemo).
REFERENCES. — Xanthias canaliculatus Rathbun, 1907: 45 (Makemo). — FOREST & GUINOT, 1962: 64 (Biogeography
"Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212 (Moorea). — GUINOT, 1985:
451 (List).
Xanthias lamarcki (H. Milne Edwards, 1834)
DISTRIBUTION. — Gambier (Tarauru-Roa, Temoe); Society (Bora Bora, Moorea, Raiatea, Tahiti); Tuamotu (Fakarava,
Hikueru, Makatea, Makemo, Marutea South, Mataiva, Raroia, Takapoto, Tikehau).
REFERENCES. — Xanthodes granoso-manus Dana, 1852a: 75; 1852b: 175; 1855, pl. 8, fig. 10a-c (Society, Tuamotu).
— Xanthias lamarckii - NoBILI, 1907: 393 (Hikueru, "Timoe" = Temoe). — RATHBUN, 1907: 44 (Bora Bora, Fakarava,
Makemo, Tahiti). — SENDLER, 1923: 37 (Makatea). — BOONE, 1934: 131, pl. 70 (Raiatea, Tahiti). — HOLTHUIS, 1953:
26 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST & GUINOT, 1961: 70, fig. 63, 66a-b (Hikueru, Marutea, Tahiti,
"Taraourou-roa” = Tarauru-Roa); 1962: 64 (Biogeography "Tahiti-Tuamotu"), — PEYROT-CLAUSADE, 1977a, annex of
the species: 27; 1977b: 212 (Moorea); 1989: 113 (Tikehau). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva,
Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — SERENE, 1984: 195, fig. 112, pl. 27b (Syn.). — DELESALLE, 1985: 305
(Mataiva). — GUINOT, 1985: 451 (List).
Xanthias latifrons (De Man, 1888)
DISTRIBUTION. — Society (Moorea, Tahiti).
REFERENCES. — Xanthias latifrons - FOREST & GUINOT, 1961: 70, fig. 67a-b (Tahiti); 1962: 64 (Biogeography "Tahiti-
Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a; 1987: 9 (Moorea). — GUINOT, 1985: 451 (List).
Xanthias nitidulus (Dana, 1852)
DISTRIBUTION. — Tuamotu (Marutea South).
REFERENCES. — Xanthodes nitidulus Dana, 1852a: 76; 1852b: 177; 1855, pl. 8, fig. 1la-c (Tuamotu). — Xanthias
nitidulus - NOBILI, 1907: 392 (Marutea South).
i i a Ee EEE EET ——EEE— eee
51
REMARK. — FOREST & GUINOT (1961: 72) have placed these two references under Xanthias tetraodon, with a ? and
this remark "Nous nous abstiendrons pour l'instant de tirer des conclusions définitives, mais il est probable que
lorsqu’on disposera d'une série de Xanthia tetraodon de diverses tailles, l'on sera amené a désigner cette espéce sous le
nom de Xanthias nitidilus (Dana)".
Xanthias punctatus (H. Milne Edwards, 1834)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Xanthias punctatus - FOREST & GUINOT, 1961: 68, fig. 61, 65a-b (Tahiti); 1962: 64 (Biogeography
"Tahiti-Tuamotu"). — GUINOT, 1985: 451 (List).
Xanthias tetraodon (Heller, 1865)
DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti); Tuamotu (Hao, Hikueru, Makatea, Marutea South?, Mataiva,
Raroia, Takapoto).
REFERENCES. — Eudora tetraodon Heller, 1865: 14, pl. 2, fig. 3 (Auckland = ? Tahiti in FOREST & GUINOT, 1961). —
Xantho (Eudora) tetraodon - NoBILI, 1907: 389 (Hao). — Juxtaxanthias tetraodon - WARD, 1942: 92 (Mangareva).
— HOLTHUIS, 1953: 22 (Raroia). — MORRISON, 1954: 16 (Raroia). — Xanthias tetraodon - FOREST & GUINOT, 1961:
72, fig. 61, 68a-c, 69bis (Hikueru, Tahiti); 1962: 64 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (Raroia);
1985: 451 (List). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289
(Mataiva).
REMARK. — ODHNER (1925) and FOREST & GUINOT (1961) consider that the type locality, Auckland, mentioned by
HELLER (1865) for the description of Eudora tetraodon, is a mistake, and that it could very likely be Tahiti. Moreover,
FOREST & GUINOT (1961) mention that this species could be a synonym of Xanthias nitidilus (Dana, 1852) (cf. above).
SUBFAMILY PANOPEINAE
Panopeus pacificus Edmondson, 1931
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Panopeus pacificus - FOREST & GUINOT, 1961: 116, fig. 102, 103a-b, 104, 10Sa-b, pl. 4, fig. 3
(Tahiti); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List).
SUBFAMILY KRAUSSIINAE
Palapedia marquesas (Seréne, 1972)
DISTRIBUTION. — Tuamotu (Anaa).
REFERENCES. — Kraussia marquesas Seréne, 1972: 53, fig. 14-15, 23g, k (Anaa). — GUINOT, 1985: 449 (List). —
Palapedia marquesas - NG, 1993: 141 (subfamily nov. and gen. nov.).
Palapedia rastripes (Miller, 1887)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Kraussia rastripes - MONTEFORTE, 1984: 171, annex 1, tab. a (Tahiti). — GUINOT, 1985: 449 (List). —
Palapedia rastripes - NG, 1993: 141 (subfamily nov. and gen. nov.).
SUBFAMILY ETISINAE
Etisus bifrontalis (Edmonson, 1935)
DISTRIBUTION. — Tuamotu (Hikueru).
52
REFERENCES. — Etisodes electra - NOBILI, 1907: 390 (Hikueru) pro parte not Etisodes electra (Herbst, 1801) = Etisus
aff. bifrontalis fide GUINOT (1964: 56, 61; cf. Remark under E. electra). — Efisus aff. bifrontalis - GUINOT, 1964: 61
(Hikueru); 1985: 450 (List) = E. bifrontalis fide SERENE (1984: 230).
Etisus anaglyptus H. Milne Edwards, 1834
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Efisus anaglyptus - MONTEFORTE, 1984: 170, annex 1, tab. a; 1987: 9 (Moorea). — GUINOT, 1985:
450 (List).
Etisus demani Odhner, 1925
DISTRIBUTION. — Society (Tahiti); Tuamotu (Hikueru?).
REFERENCES. — Efisus demani - MONTEFORTE, 1984: 170, annex 1, tab. a (Tahiti). — SERENE, 1984: 227, fig. 140,
143a, pl. 31f (Hikueru?; cf. Remark). — GUINOT, 1985: 450 (List).
REMARK. — The location "Hikueru" in SERENE (1984: 227) is only mentioned in the observations. It is not indicated
under the material examined, and was not retrieved in the references cited by SERENE. It could be an erroneous reading
in GUINOT (1964: 59), where "Hikueru" is cited under Etisus frontalis Dana, just beneath E. demani Odhner.
Etisus dentatus (Herbst, 1785)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Taiaro).
REFERENCES. — Efisus dentatus - BOONE, 1934: 119, pl. 62-63 (Tahiti). — FOREST & GUINOT, 1961: 86, fig. 80a-b
(Tahiti); 1962: 64 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, annex 1, tab. a (Moorea, Tahiti);
1987: 9 (Moorea). — GUINOT, 1985: 450 (List). — POUPIN, 1994a: 41, fig. 37, pl. 4h (Tahiti, Taiaro).
Etisus electra (Herbst, 1801)
DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti); Tuamotu (Fakarava, Hikueru, Marutea South).
REFERENCES. — Efisus rugosus Jacquinot, 1852, pl. 4, fig. 2. — JacQUINOT & Lucas, 1853: 33 (Mangareva). —
Etisodes electra - NOBILI, 1907: 390 (Hikueru, "Rikitea" = Mangareva, Marutea South) pro parte cf. Remark. —
RATHBUN, 1907: 42 (Fakarava). — FOREST & GUINOT, 1961: 89, fig. 82a-b (Tahiti); 1962: 66 (Biogeography "Tahiti-
Tuamotu"). — GUINOT, 1985: 450 (List). — Etisus electra - MONTEFORTE, 1984: 170, annex 1, tab. a (Tahiti). —
SERENE, 1984: 228 (Syn.).
REMARK. — GUINOT (1964: 54) indicates that the material from Hikueru, attributed by NOBILI to E. electra, includes in
fact three species: E. electra, Etisus frontalis (Dana, 1852), and E. aff. bifrontalis Edmonson, 1935).
Etisus frontalis (Dana, 1852)
DISTRIBUTION. — Society (Moorea); Tuamotu (Hikueru).
REFERENCES. — Etisodes electra - NOBILI, 1907: 390 (Hikueru) pro parte not Etisus electra (Herbst, 1801) = E.
frontalis fide GUINOT (1964: 54; cf. Remark under E. electra). — Etisus frontalis - GUINOT, 1964: 54 (Hikueru). —
ODINETZ, 1983: 209 (Moorea). — SERENE, 1984: 229, fig. 139, pl. 3le (Syn.). — GUINOT: 1985: 450 (List). — Etisodes
frontalis - GUINOT, 1985: 450 (List; cf. Remark).
REMARK. — In her list, GUINOT (1985: 450) mentions the two following species: "Etisodes frontalis (Dana, 1852) and
Etisus frontalis Dana, 1852". It is obviously a mistake for a single species, described under Etisodes frontalis by DANA
(1852b: 187).
Etisus laevimanus Randall, 1839
DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora); Tuamotu (Mataiva).
REFERENCES. — Etisus macrodactylus - JACQUINOT, 1852, pl. 9, fig. 2 (Mangareva). — JACQUINOT & Lucas, 1853: 30
(Mangareva). — Etisus laevimanus - NoBILI, 1907: 390 (""Rikitea" = Mangareva). — RATHBUN, 1907: 42 (Bora Bora).
— Forest & GUINOT, 1961: 88 (Mangareva); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170,
a Te RTE rs NO
53
annex 1, tab. a (Mataiva). — SERENE, 1984: 225 (Syn.). — GUINOT, 1985: 450 (List). — DELESALLE, 1985: 289
(Mataiva). — POUPIN, 1994a: 42, fig. 38, pl. 5a (Mangareva) - SYNONYMS - Etisus macrodactylus Bianconi, 1851.
Etisus punctatus Jacquinot, 1852
DISTRIBUTION. — Gambier (Mangareva).
REFERENCES. — Etisus punctatus Jacquinot, 1852, pl. 3, fig. 5. — JACQUINOT & LUCAS, 1853: 31 (Mangareva).
REMARK. — This Etisus has been figured by JACQUINOT, then described by JACQUINOT & LUCAS with this commentary
"Cette espéce n'ayant pas été déposée au Muséum, c'est d'aprés la figure qui en a été donnée par MM. HOMBRON et
JACQUINOT que nous avons fait cette description". According to JACQUINOT & LUCAS, Etisus punctatus is related to E.
macrodactylus Bianconi, 1851 (= E. laevimanus Randall, 1851) and to E. anaglyptus H. Milne Edwards, 1834.
Etisus splendidus Rathbun, 1906
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru, Mataiva?, Raroia, Taiaro, Takapoto).
REFERENCES. — Etisus (Etisodes) splendidus - HOLTHUIS, 1953: 21 (Raroia). — MORRISON, 1954: 16 (Raroia). —
Etisus splendidus - FOREST & GUINOT, 1961: 88, fig. 8la-c (Hikueru); 1962: 66 (Biogeography "Tahiti-Tuamotu"). —
GUINOT, 1966a: 48 (Raroia); 1985: 450 (List). — MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva?, Moorea, Tahiti,
Takapoto?). — SALVAT, 1986b: 72, photograph (French Polynesia). — BONVALLOT et al., 1994: 145 (Tuamotu). —
POUPIN, 1994a: 43, fig. 39, pl. 5b (Taiaro, Takapoto).
SUBFAMILY CHLORODIINAE
Chlorodiella barbata (Borradaile, 1900)
DISTRIBUTION. — Gambier (Mangareva); Society (Moorea, Tahiti); Tuamotu (Mataiva, Marutea South, Takapoto,
Tikehau).
REFERENCES. — Chlorodiella barbata - FOREST & GUINOT, 1961: 96, fig. 93-94, 99a-b, 100 (Mangareva, Marutea
South, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu”). — PEYROT-CLAUSADE, 1977a, annex of the species: 26;
1977b: 213; 1985: 462 (Moorea); 1989: 112 (Tikehau). — NAIM, 1980a, annex 1, tab. 3; 1980b: 550 (Moorea). —
THOMASSIN et al., 1982: 394 (Moorea). — ODINETZ, 1983: 97 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a
(Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GALZIN & POINTIER,
1985: 100 (Moorea). — GUINOT, 1985: 450 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea
and/or Tahiti). — SALVAT & RICHARD, 1985: 360 (Takapoto).
Chlorodiella cytherea (Dana, 1852)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru, Mataiva, Raraka, Takapoto, Tikehau).
REFERENCES. — Chlorodius cytherea Dana 1852a: 79; 1852b: 213; 1855, pl. 12, fig. 2a-c (Raraka, Tahiti). —
Chlorodiella cytherea - FOREST & GUINOT, 1961: 95, fig. 90-92, 98a-b (Hikueru, Tahiti); 1962: 66 (Biogeography
"Tahiti-Tuamotu”). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 213 (Moorea); 1989: 112 (Tikehau).
— NAIM, 1980a, annex 1, tab. 3 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva, Moorea, Tahiti,
Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 450 (List).
Chlorodiella laevissima (Dana, 1852)
DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Tahiti); Tuamotu
(Fakarava, Hao, Makatea, Makemo, Mataiva, Rangiroa, Takapoto, Tikehau) - Littoral to sublittoral.
REFERENCES. — Chlorodius laevissimus - NOBILI, 1907: 393 (‘‘Rikitea, Gatavake" = Mangareva, "Ohura" = Hao). —
Chlorodiella laevissima - RATHBUN, 1907: 46 (Fakarava, Makemo, Rangiroa, Tahiti; 46m). — FOREST & GUINOT,
1961: 95, fig. 95-96, 101a-b (Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex
of the species: 26; 1977b: 213; 1985: 462 (Moorea); 1989: 111, 114 (Moorea, Tikehau). — ODINETZ, 1983: 209
(Moorea, Tahiti). — MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9
54
(Moorea). — SERENE, 1984: 260, fig. 171-172, pl. 36d-e (Syn.). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985:
450 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti). — Chlorodiella laevissimus
- BOONE, 1934: 138, pl. 72 (Nuku Hiva, Tahiti).
Chlorodiella nigra (Forskal, 1775)
DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Hao, Makatea,
Makemo, Marutea South, Mataiva, Nukutipipi, Rangiroa, Raraka, Takapoto, Tikehau).
REFERENCES. — Chlorodius niger - HELLER, 1865: 18 (Tahiti). — STIMPSON, 1858a: 33; 1907: 50 (Tahiti). — NosILI,
1907: 393 (""Gatavake" = Mangareva, Hao, Marutea, "Tikahan" = Tikehau). — Chlorodiella niger - RATHBUN, 1907: 46
(Bora Bora, Fakarava, Makemo, Rangiroa). — SENDLER, 1923: 38 (Makatea). — SEURAT, 1934: 59 ("Gatavake" =
Mangareva, Hao, Marutea South, "Tikahau" = Tikehau). — Chlorodiella nigra - FOREST & GUINOT, 1961: 95, fig. 87-
89, 97a-b (Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26;
1977b: 213 (Moorea). — KROPP & BIRKELAND, 1981: 630, tab. 5 (Moorea, Takapoto). — THOMASSIN et al., 1982: 394
(Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a, photograph p. 136a (Mataiva, Moorea, Tahiti, Takapoto); 1987:
9 (Moorea). — SERENE, 1984: 258, fig. 168, pl. 36b (Syn.). — DELESALLE, 1985: 288 (Mataiva). — GUINOT, 1985: 450
(List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti, Takapoto). — MERSCHARDT-
SALVAT, 1991: 89 (Nukutipipi).
Garthiella aberrans (Rathbun, 1906)
DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral.
REFERENCES. — Pilodius aberrans - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212 (Moorea); 1989:
111, 115 (Moorea, Tikehau; 30m). — GUINOT, 1985: 451 (List). — Garthiella aberrans - TITGEN, 1986: 57, fig. 1-2
(Syn.).
Liocarpilodes armiger (Nobili, 1906)
DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral.
REFERENCES. — Liocarpilodes armiger - PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 213 (Moorea);
1989: 111, 115 (Moorea, Tikehau; 30m). — GUINOT, 1985: 450, with a ? (List).
Liocarpilodes harmsi (Balss, 1934)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Pilodius harmsi - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 213 (Moorea). —
Liocarpilodes harmsi - SERENE, 1984: 264 (Syn.).
Liocarpilodes integerrimus (Dana, 1852)
DISTRIBUTION. — Gambier (Mangareva); Society (Moorea, Tahiti); Tuamotu (Fakarava, Mataiva, Takapoto, Tikehau).
REFERENCES. — Actumnus integerrimus - RATHBUN, 1907: 56, pl. 1, fig. 12, pl. 8, fig. 3, 3a-b (Fakarava, Tahiti). —
Pilumnus margaritatus - NOBILI, 1907: 398 ("Rikitea" = Mangareva) not Pilumnus margaritatus Ortmann, 1893 = L.
integerrimus fide GUINOT (1964: 63). — Liocarpilodes integerrimus - FOREST & GUINOT, 1962: 64 (Biogeography
"Tahiti-Tuamotu"). — GUINOT, 1964: 63, fig. 36a-b ("Rikitea" = Mangareva). — PEYROT-CLAUSADE, 1977a, annex of
the species: 26; 1977b: 213; 1985: 462 (Moorea); 1989: 111, 115 (Moorea, Tikehau). — MONTEFORTE, 1984: 171,
annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). —
GUINOT, 1985: 450 (List).
Phymodius granulosus (De Man, 1888)
DISTRIBUTION. — Society (Moorea and/or Tahiti).
REFERENCES. — Phymodius granulosus - GUINOT, 1985: 451, with a ? (List). — ODINETZ-COLLART & RICHER DE
FORGES, 1985: 201 (Moorea and/or Tahiti).
55
REMARK. — ODINETZ does not mention this material in her thesis (1983), but it is recorded in the Museum of Paris
(MNHN B17071, coll. ODINETZ 1981 "Moorea, Tahiti, Takapoto, associé au Pocillopora damicornis et P. elegans", det.
GUINOT).
Phymodius monticulosus (Dana, 1852)
DISTRIBUTION. — Gambier (Mangareva); Society (Moorea, Tahiti); Tuamotu (Fakahina, Fakarava, Mataiva, Marutea
South, Takapoto).
REFERENCES. — Chlorodius monticulosus Dana, 1852a: 79; 1852b: 206; 1855, pl. 11, fig. 9a (Tahiti). — STIMPSON,
1858a: 31; 1907: 50 (Tahiti). — Chlorodius Dehaanii - HELLER, 1865: 19 (Tahiti) pro parte, some sp. attributed to
Phymodius ungulatus in FOREST & GUINOT (1961: 106, 114), not C. Dehaani, synonym of Phymodius granulatus
(Targioni Tozzetti, 1877) in SERENE (1984: 250). — Cyclodius ornatus - NOBILI, 1907: 397 (Fakahina, Marutea South).
— RATHBUN, 1907: 51, pl. 5, fig. 5, pl. 7, fig. 8 (Fakarava, Tahiti). — Phymodius monticulosus - FOREST & GUINOT,
1961: 106, pl. 10, fig. 1-6 (Fakahina, Marutea South, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — NAIM,
1980a, annex 1, tab. 3 (Moorea). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto);
1987: 9 (Moorea). — GUINOT, 1985: 451 (List). — POUPIN, 1994a: 44, fig. 40, pl. Sc (Fakahina, Mangareva) -
SYNONYMS - Cyclodius ornatus Dana, 1852.
Phymodius nitidus (Dana, 1852)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu ( Kaukura, Tikehau).
REFERENCES. — Pilodius nitidus - NOBILI, 1907: 393 (Kaukura). — Chlorodopsis scabricula - RATHBUN, 1907: 50, pl.
1, fig. 3, pl. 9, fig. 5 (Tahiti) not Pilodius scabriculus (Dana, 1852) = Phymodius nitidus fide FOREST & GUINOT (1961:
114). — Phymodius nitidus - FOREST & GUINOT, 1961: 114, pl. 15, fig. 1-4 (Kaukura, Tahiti); 1962: 68 (Biogeography
"Tahiti-Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a; 1987: 9 (Moorea). — GUINOT, 1985: 451 (List). —
PEYROT-CLAUSADE, 1989: 113 (Tikehau).
Phymodius ungulatus (H. Milne Edwards, 1834)
DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Raiatea, Tahiti); Tuamotu (Fakahina, Fakarava,
Hikueru, Makemo, Mataiva, Rangiroa, Raroia, Takapoto, Tikehau).
REFERENCES. — Chlorodius ungulatus - DANA, 1852b: 205; 1855, pl. 11, fig. 8a-b (Tahiti). — Chlorodius dehaanii -
HELLER, 1865: 19 (Tahiti) pro parte fide FOREST & GUINOT (1961: 110), not C. Dehaani, synonym of Phymodius
granulatus (Targioni Tozzetti, 1877) in SERENE (1984: 250). — Cyclodius gracilis - NOBILI, 1907: 397 (Fakahina,
"Rikitea" = Mangareva). — Phymodius ungulatus - RATHBUN, 1907: 46, pl. 3-4 (Bora Bora, Fakarava, Makemo,
Rangiroa, Tahiti) pro parte fide FOREST & GUINOT (1961: 110; cf. Remark). — NOBILL 1907: 393 ("Rikitea" =
Mangareva). — BOONE, 1934: 140, pl. 73 (Raiatea, Tahiti) pro parte fide FOREST & GUINOT (1961: 110). — HOLTHUIs,
1953: 25 (Raroia). — FOREST & GUINOT, 1961: 110, fig. 86a-b, pl. 11, fig. 1-4, pl. 12, fig. 1-4, pl. 13, fig. 1-3, pl. 14,
fig. 1-3 (Fakahina, Hikueru, Mangareva, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1964: 74, fig.
38 (Tahiti); 1985: 451 (List). — PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 213 (Moorea); 1989: 113
(Tikehau). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). —
SERENE, 1984: 251, fig. 158, 161, pl. 35e (Tahiti; Syn.). — DELESALLE, 1985: 288 (Mataiva). — ODINETZ-COLLART &
RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti, Takapoto). — SALVAT & RICHARD, 1985: 356 (Takapoto) -
SYNONYMS - Cyclodius gracilis Dana, 1852.
REMARK. — FOREST & GUINOT (1961: 111) indicate that RATHBUN's material is only partially assignable to this species,
but without mentioning the localities referring to the real Phymodius ungulatus. Thus, for this species, some of the
islands mentioned by RATHBUN are doubtful.
Pilodius areolatus (H. Milne Edwards, 1834)
DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Hikueru, Makemo, Marutea
South, Mataiva, Raroia).
REFERENCES. — Actaeodes affinis Dana, 1852a: 78; 1852b: 197; 1855, pl. 11, fig. 3 (Tahiti, Tuamotu). — Actaea
affinis - RATHBUN, 1907: 42 (Makemo). — Chlorodopsis areolata - NOBILI, 1907: 396, pl. 2, fig. 3 (Hikueru, "Rikitea"
56
= Mangareva, Marutea South). — HOLTHUIS, 1953: 15 (Raroia). — MORRISON, 1954: 13 (Raroia). — Pilodius
areolatus - FOREST & GUINOT, 1961: 90 (Hikueru, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). —
MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva). — GUINOT, 1985: 451 (List). — CLARK & GALIL, 1993: 1125, fig.
la-g, 31a, 40a, 44b (Bora Bora, "Maharepa, Afareaitu, Temae" = Moorea; Syn.).
Pilodius flavus Rathbun, 1893
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hao, Tikehau).
REFERENCES. — Pilodius pubescens - NoBILI, 1907: 395, with a ? ("Ohura" = Hao) not Pilodius pubescens Dana, 1852
= P. flavus fide CLARK & GALIL (1993: 1130, 1146). — Pilodius flavus - FOREST & GUINOT, 1961: 95 (Tahiti); 1962:
66 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 27 (Moorea); 1989: 111, 115
(Moorea, Tikehau). — GUINOT, 1985: 451 (List). — CLARK & GALIL, 1993: 1130, fig. 4a-g, 32b, 40d, 41a (Syn.).
Pilodius paumotensis Rathbun, 1907
DISTRIBUTION. — Society (Bora Bora); Tuamotu (Fakarava, Makemo, Marutea South, Tikehau).
REFERENCES. — Chlorodopsis granulatus - NoBILI, 1906: 396 (Marutea South) not Pilodius granulatus Stimpson,
1859 = P. paumotensis fide GUINOT (1962: 238). — Pilodius paumotensis Rathbun 1907: 52, pl. 8, fig. 2, 2a-b
(Fakarava, Makemo). — FOREST & GUINOT, 1962: 66 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 451 (List).
— PEYROT-CLAUSADE, 1989: 113 (Tikehau). — CLARK & GALIL, 1993: 1143, fig. 10a-g, 35b, 43a (Bora Bora,
Fakarava, Makemo, Marutea South).
Pilodius pubescens Dana, 1852
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Pilodius pubescens - GUINOT, 1985: 451 (List; probably after NOBILI, cf. under P. flavus). — CLARK &
GALIL, 1993: 1146, fig. 12a-g, 36b, 43b (""Ternae" = Temae at Moorea, Society). — Not Pilodius pubescens - NOBILI,
1907: 395, with a ? = P. flavus in CLARK & GALIL (1993: 1130).
Pilodius pugil Dana, 1852
DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Hikueru, Makaiea,
Makemo, Mataiva, Rangiroa, Takapoto, Tikehau).
REFERENCES. — Chlorodopsis pugil - NOBILI, 1907: 395 ("Rikitea" = Mangareva). — Chlorodopsis spinipes -
RATHBUN, 1907: 50, pl. 2, fig. 5 (Bora Bora, Fakarava, Makemo, Rangiroa) not Pilodius spinipes Heller, 1861 = P.
pugil fide CLARK & GALIL (1983: 1149). — Pilumnus globosus - BOONE, 1934: 152, pl. 78 (Tahiti) not Globopilumnus
globosus (Dana, 1852) = Pilodius pugil with a ? fide SERENE & LUOM (1959: 320). — Pilodius pugil - FOREST &
GUINOT, 1961: 91 (Hikueru, Mangareva); 1962: 64 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a,
annex of the species: 27; 1977b: 213; 1985: 462 (Moorea); 1989: 113 (Tikehau). — MONTEFORTE, 1984: 171, annex 1,
tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289, 305 (Mataiva). —
GUINOT, 1985: 451 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea, Tahiti, Takapoto). —
SALVAT & RICHARD, 1985: 358, 360 (Takapoto). — CLARK & GALIL, 1993: 1149, fig. 13a-g, 37a, 43c (Hikueru,
Mangareva, "Temae, Tiahura and Afareaita" = Moorea, Tahiti, Takapoto).
Pilodius scabriculus Dana, 1852
DISTRIBUTION. — Gambier (Temoe); Society (Bora Bora, Huahine, Maiao?, Moorea, Tahaa?, Tahiti); Tuamotu
(Fakarava, Fakahina, Hao, Hikueru, Makatea, Makemo, Marutea North?, Marutea South, Mataiva, Raraka, Takapoto,
Tikehau).
REFERENCES. — Pilodius scabriculus Dana, 1852a: 80; 1852b: 220; 1855, pl. 12, fig. 9 (Raraka). — NOBILI, 1907: 394
("Fakaina" = Fakahina, Fakarava, Marutea, Marutea South, Temoe). — FOREST & GUINOT, 1961: 91, fig. 83a-b, 84,
86bis (Hikueru, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"), — PEYROT-CLAUSADE, 1977a, annex of the
species: 27; 1977b: 213 (Moorea); 1989: 113 (Tikehau). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea,
Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289, 305 (Mataiva). — GUINOT, 1985:
451 (List). — CLARK & GALIL, 1993: 1152, fig. 14a-g, 37b, 43d (Bora Bora, Fakahina, Hao, Hikueru, Huahine, Maiai =
ee ee eeeeeeeeeeeeEeEeEeEeEeEeEeEEEEEeeeeee
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Maiao?, Marutea, Moorea, Raiatea, Tahiti, Temoe, "Tickahau" = Tikehau, "Vaiorea” = Vaitoare?, Tahaa; Syn.). —
Chlorodopsis venusta Rathbun, 1907: 49, pl. 1, fig. 5 (Fakarava, Makemo).
Tweedieia laysani (Rathbun, 1906)
DISTRIBUTION. — Tuamotu (Tikehau).
REFERENCES. — Tweedieia laysani - PEYROT-CLAUSADE, 1989: 112 (Tikehau).
Tweedieia odhneri (Gordon, 1934)
DISTRIBUTION. — Society (Moorea).
REFERENCES. — Tweedieia odhneri - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977: 212 (Moorea). —
GUINOT, 1985: 451, with a ? (List).
FAMILY TRAPEZIIDAE
Jonesius triunguiculatus (Borradaile, 1902)
DISTRIBUTION. — Society (Raiatea).
REFERENCES. — Jonesius triunguiculatus - GALIL & TAKEDA, 1986: 165, fig. 1-4 (""Tetaro” = Raiatea).
Quadrella lewinsohni Galil, 1986
DISTRIBUTION. — Marquesas (Tahuata).
REFERENCES. — Quadrella sp. - MONOD, 1979: 9, fig. 1-8 (Tahuata). — Quadrella cyrenae - SERENE, 1975: 510, fig. 3-
4, pl. 1b’, e' (Tahuata; MONOD's material) not Q. cyrenae Ward, 1942, synonym of Q. maculosa Alcock, 1898 = Q.
lewinsohni nov. in GALIL (1986a: 285). — GUINOT, 1985: 452 (List). — Quadrella lewinsohni Galil, 1986a: 285, fig.
Sa-b, 6 (Marquesas; MONOD's and SERENE's material).
Quadrella maculosa Alcock, 1898
DISTRIBUTION. — Marquesas (Fatu Hiva) - Littoral to sublittoral.
REFERENCES - Quadrella maculosa - NEW MATERIAL - Coll. J. POUPIN, det. B. GALIL (Fatu Hiva; 49m).
Tetralia cinctipes Paulson, 1875
DISTRIBUTION. — Austral (Rapa) - Sublittoral.
REFERENCES. — Tetralia cinctipes - GALIL, 1986b: 97, fig. 1-3 (Rapa; 90m).
Tetralia glaberrima (Herbst, 1790)
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Raiatea, Tahiti); Tuamotu (Aratika, Fakarava, Hikueru, Makemo,
Marutea South, Tikehau).
REFERENCES. — Tetralia glaberrima - DANA, 1852b: 262; 1855, pl. 16, fig. 3a-h ("Carlshoff' = Aratika, Tahiti). —
ORTMANN, 1893b: 485 (Tahiti). — NOBILI, 1907: 404 (Marutea). — RATHBUN, 1907: 60 (Fakarava, Makemo, Tahiti).
— BOoneg, 1934: 174, pl. 89 (Raiatea, Tahiti). — SEURAT, 1934: 59 (Tahiti, Marutea South, Marquesas). — FOREST &
GUINOT, 1961: 139 (Hikueru); 1962: 70 (Biogeography "Tahiti-Tuamotu, Marquesas"). — SERENE, 1984: 281 (Syn.; cf.
Remark). — GUINOT, 1985: 452 (List). — PEYROT-CLAUSADE, 1989: 111 (Tikehau). — Trapezia serratifrons
Jacquinot, 1852, pl. 4, fig. 20-23. — JACQUINOT & LUCAS, 1853: 47 (Nuku Hiva). — Tetralia cavimana - HELLER,
1865: 26 (Tahiti) - SYNONYMS - Tetralia cavimana Heller, 1861.
REMARK. — In SERENE (1984: 281) all the above references (except BOONE, 1934, and PEYROT-CLAUSADE, 1989) can
be attributed either to T. glaberrima or toT. heterodactyla Heller, 1861. However, the specimens attributed to
cavimana, and the specimens identified to glaberrima with a figure of the male pleopod, are T. glaberrima without
hesitation.
58
Tetraloides nigrifrons (Dana, 1852)
DISTRIBUTION. — Society (Raiatea); Tuamotu (Makemo, Pukapuka, Tikehau).
REFERENCES. — Tetralia nigrifrons Dana, 1852a: 83; 1852b: 262; 1865, pl. 16, fig. 2a-d (""Honden" = Pukapuka). —
Tetraloides nigrifrons - GALIL, 1985: 72, fig. 1-3 (Makemo, Tikehau, "Taoru" = Raiatea).
Trapezia areolata Dana, 1852
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru, Takapoto).
REFERENCES. — Trapezia areolata Dana,1852a: 83; 1852b: 259; 1855, pl. 15, fig. 8a (Tahiti). — CANO, 1888: 173
(Tahiti). — ORTMANN 1893b: 485 (Tahiti; cf. Remark). — FOREST & GUINOT, 1961: 135, fig. 133 (Hikueru). —
ODINETZ, 1983: 31 (French Polynesia). — ODINETZ, 1984a: 443, fig. 3c, 4c (Moorea, Tahiti, Takapoto). — GALIL &
LEWINSOMN, 1985a: 286, fig. 1, 3-4 (Tahiti). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES,
1985: 201 (Moorea, Tahiti, Takapoto). — ? Trapezia ferruginea areolata - SEURAT, 1934: 59 (Tahiti). — Trapezia
reticulata - KROPP & BIRKELAND, 1981: 629 (Moorea, Takapoto). — ODINETZ, 1983: 31, 205 photograph 3 (Moorea,
Tahiti, Takapoto). — GUINOT, 1985: 452 (List; after ODINETZ, 1983) - All, not Trapezia reticulata Stimpson, 1858,
synonym of T. septata Dana, 1852 =T. areolata fide ODINETZ (1984a: 443; does not mention KRopp & BIRKELAND, but
it is probably the same material).
REMARK. — In GALIL & LEWINSOHN (1985a), ORTMANN's (1893b) work, in which T. aerolata is recorded from Tahiti,
New Guinea, and Palau, is cited at the same time under T. aerolata and T. septata. It is probable that the two Tahitian
specimens belong to T. areolata, and the other to T. septata (opt. cit.: 291).
Trapezia bella Dana, 1852
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Aratika, Hikueru, Mataiva, Takapoto).
REFERENCES. — Trapezia bella Dana, 1852a: 83; 1852b: 254; 1855, pl. 15, fig. 2 ("Carlshoff' = Aratika). — FOREST &
GUINOT, 1961: 133, fig. 129-130, 135a-b (Hikueru); 1962: 70 (Biogeography "Tahiti-Tuamotu"). — KrRopp &
BIRKELAND, 1981: 629 (Moorea, Takapoto). — ODINETZ, 1983: 206 (Tahiti, Takapoto). — SERENE, 1984: 278 (Syn.).
— DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985:
201 (Tahiti?, Takapoto). — Trapezia digitalis bella - RATHBUN, 1907: 59 (Tahiti). — Not Trapezia bella Dana -
NoBILI, 1907: 403 (Hao). — SEURAT, 1934: 59 (Hao; NOBILI's material) = Trapezia speciosa fide SERENE (1984: 278).
Trapezia cymodoce (Herbst, 1799)
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Makemo,
Rangiroa, Takapoto).
REFERENCES. — Trapezia dentata - DANA, 1852b: 258; 1855, pl. 15, fig. 6a-d (Tahiti). — Trapezia hirtipes Jacquinot,
1852, pl. 4, fig. 14-16 (Nuku Hiva). — JAcQuINOT & Lucas, 1853: 44 (Nuku Hiva). — Trapezia cymodoce dentata -
RATHBUN, 1907: 58 (Bora Bora, Fakarava, Makemo, Rangiroa; cf. Remark). — Trapezia cymodoce ferruginea -
RATHBUN, 1907: 58 (Bora Bora, Fakarava, Makemo, Rangiroa; cf. Remark). — Trapezia ferruginea dentata - SEURAT,
1934: 59 (Tahiti). — Trapezia cymodoce - ORTMANN, 1897b: 203 (Tahiti). — SEURAT, 1934: 59 (Tahiti, Marquesas).
— ODINETZ, 1983: 205, photograph 2 (Moorea, Takapoto, Tahiti); 1984a: 432, fig. 1-2 (Moorea, Tahiti, Takapoto; pro
parte, some specimens would belong to T. ferruginea, cf. Remark under that species). — GUINOT, 1985: 452 (List). —
ODINETZ-COLLART & RICHER DE FORGES, 1985: 201(Moorea?, Tahiti, Takapoto). — GALIL & CLARK, 1990: 378 (Syn.).
— Trapezia cymodoce sp.1 - KRopP & BIRKELAND, 1981: 629 (Moorea, Takapoto) fide distinction in ODINETZ (1984b:
124). — ODINETZ, 1984b: 125 (Moorea, Tahiti, Takapoto). — Not Trapezia cymodoce - DANA, 1852b: 257; 1855, pl.
15, fig. 5a-i (Tahiti). — SENDLER, 1923: 39 (Tahiti) - The two references = T. ferruginea Latreille 1825 fide GALIL &
CLARK (1990: 378) - SYNONYMS - Trapezia dentata (Macleay, 1838).
REMARK. — The material attributed to T. cymodoce dentata by RATHBUN (1907) has been partially re-examined by
GALIL & CLARK (1990; sp. of Ellice islands "Funafuti reef") and belongs to T. cymodoce. Arbitrarily we also attribute to
this species the material from French Polynesia. Concerning the T. cymodoce ferruginea also recorded by this author,
from French Polynesia, Ellice, and Easter island, they would belong in part to that species (?Bora Bora, Fakarava,
Makemo, Rangiroa), in part to T. guttata Riippell (2 sp. from "Mohican reef" at Rangiroa) (cf. GALIL & CLARK, 1990:
380-382), and in part to T. punctimanus (sp. from Easter island; cf. ODINETZ 1984a: 446).
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59
Trapezia digitalis Latreille, 1825
DISTRIBUTION. — Marquesas (Nuku Hiva); Tuamotu (Makatea, Takapoto).
REFERENCES. — Trapezia fusca Jacquinot, 1852, pl. 4, fig. 17-19. — JACQUINOT & Lucas, 1853: 45 (Nuku Hiva). —
Trapezia digitalis - SENDLER, 1923: 40 (Makatea). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu,
Marquesas”). — ODINETZ, 1983: 31, 206 (Takapoto; cf. Remark). — SERENE, 1984: 277 (Syn.). — GUINOT, 1985: 452
(List).
REMARK. — Although ODINETZ (1983: 31) has indicated that her material was collected at Guam, one specimen in table
9 (p. 206) is recorded from Takapoto.
Trapezia ferruginea Latreille, 1825
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Raiatea, Tahiti); Tuamotu (Takapoto).
REFERENCES. — Trapezia ferruginea - DANA, 1852b: 260; 1855, pl. 16, fig. 1a-b (Tahiti, pro parte, see under T.
guttata; Samoan specimens are T. serenei fide ODINETZ, 1984a: 440, 442). — BOONE, 1934: 171, pl. 88 (Bora Bora,
Nuku Hiva, Raiatea). — SEURAT, 1934: 59 (Tahiti, Marquesas). — FOREST & GUINOT, 1961: 136, fig. 137a-b (Tahiti).
— Kropp & BIRKELAND, 1981: 629 (Moorea). — SERENE, 1984: 273 (Syn.). — GUINOT, 1985: 452 (List). — PEYROT-
CLAUSADE, 1989: 115 (Moorea). — Trapezia cymodoce - DANA, 1852b: 257; 1855, pl. 15, fig. 5a-i (Tahiti). —
SENDLER, 1923: 39 (Tahiti) - The two references not Trapezia cymodoce (Herbst, 1799) = T. ferruginea fide GALIL &
CLARK (1990: 380). — ODINETZ, 1984a: 432 (Moorea, Tahiti, Takapoto; pro parte, cf. Remark). — Trapezia miniata
Jacquinot, 1852, pl. 4, fig. 10-13. — Jacquinot & Lucas, 1853: 43 (Nuku Hiva). — Not Trapezia cymodoce
ferruginea - RATHBUN, 1907: 58 (cf. Remark under T. cymodoce).
REMARK. — Trapezia ferruginea Latreille, 1825 was proposed as a synonym of T. cymodoce (Herbst, 1799) in ODINETZ
(1984a), but this proposition was not followed by GALIL & CLARK (1990).
Trapezia flavopunctata Eydoux & Souleyet, 1842
DISTRIBUTION. — Society (Moorea,Tahiti); Tuamotu (Hikueru).
REFERENCES. — Trapezia flavopunctata - ORTMANN, 1893b: 485 (Tahiti). — FOREST & GUINOT, 1961: 136, fig. 138a-b
(Hikueru). — ODINETZ, 1983: 34, 205 (Tahiti; p. 205 = "T. flavomaculata" sic). — GALIL & LEWINSOHN, 1985b: 210
("Papetoai bay" = Moorea, Tahiti). — GUINOT, 1985: 452 (List). — Trapezia rufopunctata flavopunctata - SEURAT,
1934: 59 (Tahiti).
Trapezia formosa Smith, 1869
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Mataiva, Takapoto).
REFERENCES. — Trapezia formosa - KROPP & BIRKELAND, 1981: 629 (Takapoto). — ODINETZ, 1983: 206, photograph
4 (Moorea, Takapoto, Tahiti). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART
& RICHER DE FoRGES, 1985: 201 (Moorea?, Takapoto, Tahiti).
REMARK. — According to P. CASTRO (personal communication), who has re-examinated the material of these
references, it could rather belongs to a new species.
Trapezia guttata Riippell, 1830
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru, Rangiroa, Takapoto, Tikehau).
REFERENCES. — Trapezia guttata - HELLER, 1865: 25 (Tahiti). — FOREST & GUINOT, 1961: 136, fig. 134, 139a-b
(Hikueru, Tahiti); 1962: 70 (Biogeography "Tahiti-Tuamotu"). — ODINETZ, 1983: 205, photograph 8 (Moorea, Tahiti,
Takapoto); 1984a: 442 (Moorea, Tahiti). — GuINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES,
1985: 201 (Moorea?, Tahiti, Takapoto). — PEYROT-CLAUSADE, 1989: 111 (Tikehau). — GALIL & CLARK, 1990: 381
(Syn.). — Trapezia cymodoce ferruginea - RATHBUN, 1907: 58 (only some specimens from Rangiroa) not T.
ferruginea Latreille, 1825 = T. guttata fide GALIL & CLARK (1990: 381; cf. Remark under T. cymodoce). — Trapezia
ferruginea - DANA, 1852b: 260; 1865, pl. 16, fig. 1b (Tahiti) pro parte not T. ferruginea Latreille, 1825 = T. guttata
fide GALIL & CLARK (1990: 381, 382). — Trapezia ferruginea guttata - SEURAT, 1934: 59 (Tahiti). — Trapezia
60
davaoensis - KROpP & BIRKELAND, 1981: 629 (Moorea). — GUINOT, 1985: 452 (List) - SYNONYMS - Trapezia
davaoensis Ward, 1941.
Trapezia punctimanus Odinetz, 1984
DISTRIBUTION. — Society (Tahiti); Tuamotu (Takapoto).
REFERENCES. — Trapezia punctimanus Odinetz, 1983: 35, 206 photograph 7 (Thesis; French Polynesia); 1984a: 445,
fig. 3e, 4e (Tahiti, Takapoto). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201
(Tahiti, Takapoto).
Trapezia rufopunctata (Herbst, 1801)
DISTRIBUTION. — Society (Moorea, Raiatea, Tahiti); Tuamotu (Makemo, Takapoto, Tikehau).
REFERENCES. — Trapezia rufo-punctata - DANA, 1852b: 255; 1855, pl. 15, fig. 3a-b (Tahiti). — Trapezia rufopunctata
- RATHBUN, 1907: 57 (Makemo). — BOONE, 1934: 166, pl. 86 (Raiatea). — SEURAT, 1934: 59 (Tahiti, but not the
Marquesas certainly cited after JACQUINOT & LUCAS; see below). — ODINETZ, 1983: 34 (Moorea, Tahiti, Takapoto). —
GUINOT, 1985: 452 (List). — PEYROT-CLAUSADE, 1989: 112 (Tikehau). — Not Trapezia rufo-punctata - JACQUINOT,
1852, pl 4, fig. 8-9. — JACQUINOT & LucAs, 1853: 41 (Nuku Hiva) = T. tigrina Eydoux & Souleyet, 1842 fide GALL &
LEWINSOHN (1985b: 166).
Trapezia septata Dana, 1852
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Trapezia ferruginea areolata - SENDLER, 1923: 40 (Tahiti) not T. areolata Dana, 1852 = T. septata
fide GALIL & LEWINSOHN (1985a: 288; cf. Remark).
REMARK. — Although SENDLER is cited under that species in GALIL & LEWINSOMN, these authors have not examined
Polynesian material. They only mention that (p. 291) "T. septata seems to be more widely distributed and more
common than T. areolata. Thus, specimens identified as T. aerolata, with the exception of those mention by CANO
(1888), ORTMANN (1893) (part) and FOREST & GUINOT (1961), should rightly be name T. septata".
Trapezia serenei Odinetz, 1984
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Mataiva, Takapoto).
REFERENCES. — Trapezia serenei Odinetz, 1983: 34, 206, photograph 6; 1984a: 440, fig. 3b, 4b (Mataiva, Moorea,
Tahiti, Takapoto). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea?,
Tahiti, Takapoto). — SALVAT & RICHARD, 1985: 344 (Takapoto). — Trapezia cymodoce sp. 2 - KROPP & BIRKELAND,
1981: 629 (Moorea, Takapoto). — ODINETZ, 1984b: 125 (Moorea, Tahiti, Takapoto) - Fide distinction in ODINETZ
(1984b: 124).
Trapezia speciosa Dana, 1852
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Aratika, Fakarava, Hao, Hikueru, Makemo, Marutea South,
Mataiva, Takapoto).
REFERENCES. — Trapezia speciosa Dana, 1852a: 83; 1852b: 253; 1855, pl. 15, fig. 1 ("Carlshoff' = Aratika). —
NosILI, 1907: 403 (Marutea). — SEURAT, 1934: 59 (Marutea South). — FoREST & GUINOT, 1961: 133, fig. 131-132,
136a-b (Hikueru); 1962: 70 (Biogeography "Tahiti-Tuamotu"). — KROpP & BIRKELAND, 1981: 629 (Moorea,
Takapoto). — ODINETZ, 1983: 205 (Moorea, Tahiti, Takapoto). — SERENE, 1984: 278 (Syn.). — DELESALLE, 1985: 289
(Mataiva). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea, Tahiti,
Takapoto). — Trapezia digitalis speciosa - RATHBUN, 1907: 59 (Fakarava, Makemo, Tahiti). — Trapezia bella -
NoBILI, 1907: 403 (Hao) not T. bella Dana, 1852 = T. speciosa fide SERENE (1984: 278).
Trapezia tigrina Eydoux & Souleyet, 1842
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Moorea, Tahiti); Tuamotu (Anaa, Makemo?, Takapoto, Tikehau).
61
REFERENCES. — Trapezia maculata - DANA, 1852b: 256; 1855, pl. 15, fig. 4 (Tahiti) not T. maculata (MacLeay, 1838)
= T. tigrina fide SERENE (1984: 275) and GALIL & LEWINSHON (1984: 166). — Trapezia rufo-punctata - JACQUINOT,
1852, pl 4, fig. 8-9. — JACQUINOT & LUCAS, 1853: 41 (Nuku Hiva) not T. rufopunctata (Herbst, 1799) =T. tigrina fide
GALIL & LEWINSOHN (1984: 166). — Trapezia rufopunctata var. maculata - ORTMANN, 1893b: 484 (Tahiti). —
Trapezia cymodoce maculata - RATHBUN, 1907: 59 (Makemo) - These two references, with a ?, not T. maculata
(MacLeay, 1838) = T tigrina fide GALIL & LEWINSOHN (1984: 167). — ? Trapezia ferruginea maculata - SEURAT,
1934: 59 (Tahiti). — Trapezia wardi - KRopP & BIRKELAND, 1981: 629 (Moorea, Takapoto). — GUINOT, 1985: 452
(List). — Trapezia tigrina - ODINETZ, 1983: 205, photograph 5 (Moorea, Tahiti, Takapoto). — GALIL & LEWINSOHN,
1984: 166, fig. 1 (Anaa, "Tikehae lagoon, Tuamotu” = Tikehau, Society; Syn.). — GUINOT, 1985: 452 (List). —
ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea?, Tahiti, Takapoto) - SYNONYMS - Trapezia wardi Seréne,
1969.
FAMILY PILUMNIDAE
Actumnus asper (Rippell, 1830)
DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Marutea South).
REFERENCES. — Actumnus bonnieri - NOBILI, 1907: 400 ("“Rikitea” = Mangareva, Marutea South). — Actumnus asper
- BALsS, 1933: 36 (Syn.). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1964: 98
(Syn.); 1985: 452 (List) - SYNONYMS - Actumnus bonnieri Nobili, 1905.
Actumnus digitalis (Rathbun, 1907)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Platypodia digitalis Rathbun, 1907: 38, pl. 1, fig. 6, pl. 9, fig. 4, 4a (Tahiti). — FOREST & GUINOT,
1962: 62 (Biogeography "Tahiti-Tuamotu"). — Actumnus digitalis - GUINOT, 1969: 225 (Syn.); 1985: 452 (List).
Actumnus globulus Heller, 1861
DISTRIBUTION. — Tuamotu (Hao, Hikueru).
REFERENCES. — Actumnus globulus - NOBILI, 1907: 400 ("Ohura" = Hao, Hikueru; cf. thereafter). — FOREST &
GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1969: 226 (writes about NOBILI's work: "par contre il
est bien possible que les "globulus" polynésiens de NOBILI (1907, p. 50 sic) soient en fait des digitalis"); 1985: 452
(List).
Actumnus obesus Dana, 1852
DISTRIBUTION. — Marquesas; Society (Tahiti).
REFERENCES. — Actumnus obesus - BALSS, 1933: 37 (Marquesas). — BOONE, 1934: 154, pl. 79 (Tahiti). — FOREST &
GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1985: 452 (List).
Actumnus setifer (De Haan, 1835)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Actumnus tomentosus Dana, 1852a; 1852b: 243; 1855, pl. 14, fig. 2a-c (Tahiti). —Actumnus setifer -
ORTMANN, 1893b: 474 (Syn.). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985:
452 (List).
Pilumnus merodentatus Nobili, 1906
DISTRIBUTION. — Gambier (Mangareva); Marquesas; Society (Tahiti); Tuamotu (Mataiva?).
REFERENCES. — Pilumnus merodentatus Nobili, 1906a: 263; 1907: 399 (""Rikitea” = Mangareva). — SEURAT, 1934: 60
(French Polynesia). — FOREST & GUINOT, 1961: 132, fig. 128 (Mangareva; Syn.); 1962: 70 (Biogeography "Tahiti-
Tuamotu"). — GUINOT, 1985: 452 (List). — Pilumnus longicornis merodentatus - BALSS, 1933: 16 (Mangareva,
62
NOBILI's material and also Tahiti, Marquesas) - RELEVANT MATERIAL - Pilumnus cf. merodentatus - MONTEFORTE,
1984: 171, annex 1, tab. a (Mataiva). — GUINOT, 1985: 452 (List).
Pilumnus parvulus Nobili, 1906
DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Moruroa).
REFERENCES. — Pilumnus parvulus Nobili, 1906a: 263; 1907: 398 (""Gatavake, Rikitea, Tokaerero" = Mangareva;
some sp. in pearl oyster, 25m). — SEURAT, 1934: 60 (French Polynesia). — FOREST & GUINOT, 1961: 130, fig. 126, pl.
27, fig. 1 (Mangareva); 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List). — TAKEDA &
MIYAKE, 1968: 6 (Key) - NEW MATERIAL - Coll. & det. J. POUPIN (Moruroa; Isp. in pearl oyster Pinctada
margaritifera).
Pilumnus ransoni Forest & Guinot, 1961
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Pilumnus ransoni Forest & Guinot, 1961: 130, fig. 123-124, 127, pl. 4, fig. 1-2, pl. 17, fig. 2. (Tahiti);
1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List).
Pilumnus tahitensis De Man, 1890
DISTRIBUTION. — Society (Moorea, Raiatea, Tahiti); Tuamotu (Fakarava, Marutea South, Tikehau) - Littoral to
sublittoral.
REFERENCES. — Pilumnus tahitensis de Man, 1890: 61, pl. 3, fig. 4 (Tahiti). — ORTMANN, 1893b: 437 (Tahiti). —
NoBILI, 1907: 399 (Marutea South). — RATHBUN, 1907: 56 (Fakarava). — BALSS, 1933: 25 (Tahiti). — SEURAT, 1934:
60 (Marutea South). — FoREST & GUINOT, 1961: 129, fig. 125 (Raiatea, Tahiti); 1962: 70 (Biogeography "Tahiti-
Tuamotu"). — GUINOT, 1985: 452 (List). — PEYROT-CLAUSADE, 1989: 111, 115 (Moorea, Tikehau; 30m).
FAMILY CARPILIIDAE
Carpilius convexus (Forskal, 1775)
DISTRIBUTION. — Austral (Rapa, Rurutu); Society (Moorea, Tahiti); Tuamotu (Hikueru, Makatea, Makemo, Mataiva,
Moruroa, Taiaro, Takapoto, Takaroa, Raroia) - Littoral to sublittoral.
REFERENCES. — Carpilius convexus - STIMPSON, 1858a: 32; 1907: 37 (Tahiti). — RATHBUN, 1907: 37 (Makemo). —
BOONE, 1934: 89, pl. 43-45 (Tahiti). — HOLTHUIS, 1953: 12 (Raroia). — CHABOUIS L. & F., 1954: 91, fig. 2 (French
Polynesia). — MORRISON, 1954: 16 (Raroia). — BUITENDIJK, 1960: 263 (Takaroa). — FOREST & GUINOT, 1961: 37
(Hikueru, Tahiti); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (French Polynesia); 1985: 449
(List). — CHEVALIER et al., 1968: 92, 138 (Moruroa). — MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Mataiva,
Moorea, Takapoto). — SERENE, 1984: 303, fig. 208-209 (Mataiva). — DELESALLE, 1985: 289 (Mataiva). — SALVAT &
RICHARD, 1985: 362 (Takapoto). — SALVAT, 1986b: 72 (French Polynesia). — BONVALLOT et al., 1994: 140,
photograph (Tuamotu). — POUPIN, 1994a: 45, fig. 41, pl. 5d (Hikueru, Rapa, Rurutu, Tahiti, Taiaro; up to 60m).
Carpilius maculatus (Linné, 1758)
DISTRIBUTION. — Gambier; Society (Moorea, Tahiti); Tuamotu (Fakarava, Hao, Hikueru, Makatea, Mataiva, Moorea,
Moruroa, Raraka, Raroia, Taiaro, Takapoto).
REFERENCES. — Carpilius maculatus - DANA, 1852b: 160 (Raraka). — STIMPSON, 1858a: 32; 1907: 37 (Tahiti). —
HELLER, 1865: 9 (Tahiti). — NoBILI, 1907: 386 ("Ohura" = Hao). — RATHBUN, 1907: 37 (Fakarava, Tahiti). — PESTA,
1913: 39, pl. 3, fig. 4 (Tahiti, with a ?). — BOONE, 1934: 86, pl. 39-42 (Tahiti). — SEURAT, 1934: 60 (Gambier,
Tuamotu). — HOLTHUIS, 1953: 12 (Raroia). — CHABOUIS L. & F., 1954: 91, fig. 1 (French Polynesia). — MORRISON,
1954: 16 (Raroia). — BABLET, 1972: 32, pl. 11 (French Polynesia). — FOREST & GUINOT, 1961: 37 (Hikueru, Tahiti);
1962: 60 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (French Polynesia); 1985: 450 (List). — CHEVALIER
et al., 1968: 92, 138 (Moruroa). — CROSNIER, 1984: 302, fig. 208-209, pl. 44e (Mataiva). — MONTEFORTE, 1984: 170,
ne ae eee
63
annex 1, tab. a, photograph p.140a (haut) (Makatea, Mataiva, Moorea, Tahiti, Takapoto). — DELESALLE, 1985: 289
(Mataiva). — SALVAT, 1986b: 72, photograph (French Polynesia). — PARDON, 1992: 82, photograph (Tahiti). —
BONVALLOT ef al., 1994: 141, photograph (Tuamotu). — PouPIN, 1994a: 46, fig. 42, pl. Se (Mataiva, Tahiti, Taiaro).
FAMILY MENIPPIDAE
Dacryopilumnus eremita Nobili, 1906
DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Amanu, Hao, Makatea, Marutea South, Mataiva).
REFERENCES. — Dacryopilumnus eremita Nobili, 1906a: 264; 1907: 400, pl. 2, fig. 4 (Amanu, Hao, "Rikitea" =
Mangareva; gen. and sp. nov.). — FOREST & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu"). — CROSNIER, 1984:
313, fig. 240-241, pl. 47e (Mangareva, Marutea). — MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva). — GUINOT,
1985: 450 (List).
Domecia glabra Alcock, 1899
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hao, Tikehau) - Littoral to sublittoral.
REFERENCES. — Domecia hispida - NoBILt, 1907: 404 (Hao) not Domecia hispida Eydoux & Souleyet, 1842 = D.
glabra fide FOREST & GUINOT (1961: 126). — Domecia glabra - FOREST & GUINOT, 1961: 126, fig. 115-116, 120-122,
124bis (Hao); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1962: 240, fig. 13a-b (Hao). — PEYROT-
CLAUSADE, 1977a, annex of the species: 26; 1977b: 213 (Moorea); 1989: 111 (Tikehau; 30m). — GUINOT, 1985: 450
(List).
Domecia hispida Eydoux & Souleyet, 1842
DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Moorea, Tahiti); Tuamotu (Hikueru,
Makemo, Takapoto, Tikehau).
REFERENCES. — Domecia hispida - DANA, 1852b: 251 (Tahiti). — RATHBUN, 1907: 60 (Makemo). — BOONE, 1934:
162, pl. 85 (Nuku Hiva). — FOREST & GUINOT, 1961: 126, fig. 117-119, 124bis, pl. 28, fig. 1 (Hikueru); 1962: 68
(Biogeography "Tahiti-Tuamotu, Marquesas"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 213
(Moorea); 1989: 111, 114 (Moorea, Tikehau). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — KROPP & BIRKELAND,
1981: 629, tab. 5 (Moorea, Takapoto). — ODINETZ, 1983: 205 (Moorea, Tahiti, Takapoto). — GUINOT, 1985: 450
(List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti, Takapoto). — Not Domecia
hispida - NoBILt, 1907: 404 (Hao) = Domecia glabra Alcock, 1899 fide FOREST & GUINOT (1961: 126).
Eriphia scabricula Dana, 1852
DISTRIBUTION. — Society (Tahiti); Tuamotu (Fakarava, Raroia).
REFERENCES. — Eriphia scabricula Dana, 1852a: 82; 1852b: 247; 1855, pl. 14, fig. Sa-b (Tahiti). — RATHBUN, 1907:
57 (Fakarava). — HOLTHUIS, 1953: 20 (Raroia). — MORRISON, 1954: 7 (Raroia). — FOREST & GUINOT, 1961: 123, fig.
113a-b, 114 (Tahiti); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, annex 1, tab. a (Tahiti).
— GUINOT, 1985: 450 (List).
Eriphia sebana (Shaw & Nodder, 1803)
DISTRIBUTION. — Gambier (Mangareva, Tarauru-Roa); Society (Moorea, Tahiti); Tuamotu (Fakarava, Hikueru,
Makatea, Makemo, Marutea South, Mataiva, Moruroa, Pukapuka, Rangiroa, Raroia, Taiaro, Takapoto, Tauere).
REFERENCES. — Eriphia laevimana Latr. - DANA, 1852b: 249; 1855, pl. 14, fig. 7a-c ("Honden" = Pukapuka, Society).
— CANO, 1888: 171 (Tahiti). — NoBILI, 1907: 403 (""Gatavake" = Mangareva, Tarauru-Roa, Tauere). — Eriphia
sebana - RATHBUN, 1907: 57 (Fakarava, Makatea, Makemo, "Manga Reva, Motus" = Mangareva?, Rangiroa). —
SENDLER, 1923: 39 (Makatea). — HOLTHUIS, 1953: 20 (Raroia). — MORRISON, 1954: 7 (Raroia). — FOREST & GUINOT,
1961: 122, fig. 11la-b, 112 (Hikueru, Tahiti; Syn.); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE,
1984: 170, annex 1, tab. a, photograph 140a (bas) (Makatea, Mataiva, Moorea, Tahiti, Takapoto). — DELESALLE, 1985:
64
289 (Mataiva). — GUINOT, 1985: 450 (List). — SALVAT, 1986b: 72 (French Polynesia). —POUPIN, 1994a: 50, fig. 46,
pl. 6a (Makatea, Marutea, Moruroa, Taiaro) - SYNONYMS - Eriphia laevimana Guérin, 1829-1844 in Latreille.
Globopilumnus globosus (Dana, 1852)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Ahe?, Makatea, Manihi?, Mataiva, Raraka, Raroia, Takapoto,
Tikehau).
REFERENCES. — Pilumnus globosus Dana, 1852a: 81; 1852b: 236; 1855, pl. 13, fig. 10 (Raraka, Tahiti, "Waterland’ =
Ahe and/or Manihi). — RATHBUN, 1907: 56 (Tahiti). — Pilumnus margaritatus Ortmann, 1893b: 436 (Tahiti). —
Globopilumnus globosus - BALSS, 1933: 7, pl. 1, fig. 1-2. (Tahiti) ? pro parte. — HOLTHUIS, 1953: 21 (Raroia). —
GUINOT-DUMORTIER, 1960a: 99, fig. 1-2, 5-6 (Tahiti; Syn.). — ForEST & GUINOT, 1961: 121 (Tahiti); 1962: 68
(Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti,
Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 450 (List). — PEYROT-
CLAUSADE, 1989: 113 (Tikehau). — Not Pilumnus globosus - NOBILI, 1907: 398 (""Marutea Vaitutaki") = Liocarpilodes
sp. fide GUINOT-DUMORTIER (1960a: 100). — BOonE, 1934: 152, pl. 78 (Tahiti) = Pilodius pugil with a ? fide SERENE &
LuoM (1959: 320).
Lydia annulipes (H. Milne Edwards, 1834)
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti); Tuamotu (Hikueru, Mataiva, Moruroa, Pukapuka, Raroia,
Taiaro, Takapoto).
REFERENCES. — Ruppellia annulipes - DANA, 1852b: 246; 1855, pl. 14, fig. 4a-c (Tahiti). — Lydia annulipes -
HOLTHUIS, 1953: 23 (Pukapuka, Raroia). — MORRISON, 1954: 7 (Raroia). — FOREST & GUINOT, 1961: 122, fig. 109a-b,
110 (Hikueru); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva,
Takapoto). — DELESALLE, 1985: 289 (Mataiva). —GUINOT, 1985: 450 (List). — POUPIN, 1994a: 47, fig. 43, pl. Sf
(Hikueru, Moruroa, Nuku Hiva, Taiaro). — Ruppellia granulosa A. Milne Edwards, 1867: 279 (Marquesas; new
synonymy, cf. Remark).
REMARK. — Ruppellia granulosa has been very rarely cited after its description. It is mentioned for the genus Lydia in
SAKAI (1976: 477). It has been briefly described from a single specimen: "Cette espéce se distingue de Ruppellia
annulipes par la profondeur des sillons qui limitent les lobules des régions. Ces lobules sont rugueux et granuleux. Les
pattes antérieures sont également couvertes de grosses granulations peu élevées. Les autres caractéres sont les mémes
que chez la Ruppellia annulipes". We have re-examined the type specimen (MNHN B9344, 1 do 17x25) and, after its
comparison with specimens of L. annulipes collected in the Marquesas and the Tuamotu (cf. in PouPIN, 1994a: 47), we
consider that it is a junior synonym of this species.
Ozius hawaiensis Rathbun, 1902
DISTRIBUTION. — Marquesas (Nuku Hiva); Tuamotu (Fakarava, Makemo, Rangiroa).
REFERENCES. — Ozius hawaiensis - RATHBUN, 1907: 54 (Fakarava, Makemo, Nuku Hiva, Rangiroa). — FOREST &
GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1985: 450 (List).
Ozius rugulosus Stimpson, 1858
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti).
REFERENCES. — Ozius rugulosus - HELLER, 1865: 22, pl. 3, fig. 1 (Tahiti). — PEsTA, 1913: 47 (Tahiti). — Forest &
GUINOT, 1961: 121, fig. 107a-b, 108 (Tahiti); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 450 (List).
— PouPIN, 1994a: 48, fig. 44, pl. 5g (Nuku Hiva, Tahiti).
Ozius tricarinatus Rathbun, 1907
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti).
REFERENCES. — Ozius tricarinatus Rathbun, 1907: 53, pl. 2, fig. 3 (Nuku Hiva, Tahiti). — FOREST & GUINOT, 1962: 68
(Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1985: 450 (List).
65
Ozius truncatus A. Milne Edwards, 1834
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Ozius lobatus Heller, 1865: 21, pl. 2, fig. 4 (Tahiti) fide CHILTON & BENNETT (1929: 750).
Remark. — CHILTON & BENNETT consider, with doubt, that HELLER's species is the same as Ozius truncatus, but they
do not mention "Tahiti", in the distribution of O. truncatus It is possible, as often seen in HELLER's work, that this
locality was mentioned by error.
Ozius tuberculosus H. Milne Edwards, 1834
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti).
REFERENCES. — Ozius tuberculosus - BOONE, 1934: 150, pl. 77 (Nuku Hiva). — POUPIN, 1994a: 49, fig. 45, pl. 5h
(Nuku Hiva, Tahiti).
Pseudozius caystrus (Adams & White, 1848)
DISTRIBUTION. — Gambier (Kamaka, Tarauru-Roa); Society (Moorea, Tahiti); Tuamotu (Ahe and/or Manihi, Makatea,
Mataiva, Moruroa, Raraka, Raroia, Taiaro, Takapoto).
REFERENCES. — Pseudozius planus Dana, 1852a: 81; 1852b: 233; 1855, pl. 13, fig. 6a-h (Raraka, "Waterland’ = Ahe
and/or Manihi). — EVANS, 1967: 409 ("Paumotu"; BM syntypes). — Pseudozius caystrus - NOBILI, 1907: 397
(Kamaka). — SENDLER, 1923: 38 (Makatea). — HOLTHUIS, 1953: 26 (Raroia). — MORRISON, 1954: 9 (Raroia). —
FOREST & GUINOT, 1961: 125 (Tahiti, “"Taraourou-roa" = Tarauru-Roa; Syn.); 1962: 68 (Biogeography "Tahiti-
Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto). — DELESALLE,
1985: 289 (Mataiva). — GUINOT, 1985: 450 (List). — PoUPIN, 1994a: 52, fig. 48, pl. 6c (Kamaka, Moruroa, Taiaro).
FAMILY GECARCINIDAE
Cardisoma carnifex (Herbst, 1794)
DISTRIBUTION. — Society (Bora Bora, Moorea, Raiatea, Tahiti, Tupai); Tuamotu (Ahe, Hao, Makatea, Mataiva,
Nukutipipi, Pukarua, Rangiroa, Raroia, Taiaro, Takapoto, Tauere) - Terrestrial.
REFERENCES. — Cardisoma obesum Dana, 1851: 252; 1852b: 375; 1855, pl. 24, fig. 1 (“Peacock" = Ahe). —
STIMPSON, 1858b: 100; 1907: 111 (Tahiti). — Perigrapsus excelsus Heller, 1862: 522; 1865: SO, pl. 5, fig. 1 (Tahiti).
— Cardisoma carnifex - MIERS, 1886: 220 (Tahiti). — Not, 1907: 407 (Bora Bora, Hao, Tahiti). — RATHBUN,
1907: 26 (Rangiroa, Tahiti). — SENDLER, 1923: 22 (Tahiti). — BOONE, 1934: 187, pl. 97-98 (Bora Bora). — SEURAT,
1934: 52 (Moorea, Tahiti, Tuamotu). — HOLTHUIS, 1953: 34 (Raroia). — CHABOUIS L. & F., 1954: 92, unnumbered fig.
(French Polynesia). — MORRISON, 1954: 2 (Raroia). — FOREST & GUINOT, 1961: 165 (Tahiti); 1962: 74 (Biogeography
"Tahiti-Tuamotu"). — EDMONSON, 1962: 25 (Raiatea). — GUINOT, 1966a: 48 (French Polynesia); 1985: 454 (List). —
TURKAY, 1973: 108 (Syn.). — SAKAI, 1976: 680 (Syn.). — MONTEFORTE, 1984: 174, annex 1, tab. a (Makatea,
Mataiva, Moorea, Takapoto); 1987: 6 (Moorea). — DELESALLE, 1985: 288, 295 (Mataiva). — CHARLEUX, 1986: 80,
photographs (French Polynesia). — BONVALLOT et al., 1994: 78, photograph (Tuamotu). — BAGNIS & CHRISTIAN,
1983: 110-111, photograph (Tuamotu). — MERSCHARDT-SALVAT, 1991: 40 (Nukutipipi). — SALVAT F. & B., 1992: 5
(Nukutipipi). — POUPIN, 1994a: 53, fig. 49, pl. 6e (Pukarua, Tahiti, Taiaro, Tauere, Tupuai). — Without name -
PARDON, 1992: 78, 79, photograph, double page (Tahiti, Papeete market) (det. according to the photograph).
Cardisoma hirtipes Dana, 1851
DISTRIBUTION. — Society (Tahiti) - Terrestrial.
REFERENCES. — Cardisoma hirtipes - HELLER, 1865: 35 (Tahiti). — TURKAY, 1974: 229, fig. 2, 12-13 (Tahiti). —
GUINOT, 1985: 454, with a ? (List). — POUPIN, 1994a: 53 (Text).
66
Cardisoma rotundum Quoy & Gaimard, 1834
DISTRIBUTION. — Marquesas (Nuku Hiva); Tuamotu (Tikehau) - Terrestrial.
REFERENCES. — Cardisoma rotundum - TURKAY, 1974: 234, fig. 1, 14 (""Tickahau-Atoll" = Tikehau). — POUPIN,
1994a: 54, fig. 50, pl. 6g (Nuku Hiva).
Discoplax longipes A. Milne Edwards, 1867
DISTRIBUTION. — Tuamotu (Makatea).
REFERENCES. — Discoplax longipes - SENDLER, 1923: 23, pl. 20, 1a-b (Makatea). — FOREST & GUINOT, 1962: 74
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 454 (List).
Epigrapsus politus Heller, 1862
DISTRIBUTION. — Society (Tahiti); Tuamotu (Hikueru, Taiaro).
REFERENCES. — Epigrapsus politus Heller, 1862: 522 (Tahiti). — HOLTHUIS, 1953: 34 ("Taravao" = Tahiti). — FOREST
& GUINOT, 1961: 162, fig. 176a-b (Hikueru; Syn.); 1962: 74 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984:
172, annex 1, tab. a (Tahiti). — GUINOT, 1985: 454 (List). — POUPIN, 1994a: 55, fig. 51, pl. 6d (Hikueru). —
Nectograpsus politus Heller, 1865: 57, pl. 5, fig. 3 (Tahiti). — Not Epigrapsus politus - NOBILI, 1907: 407 (Hikueru) =
Cyclograpsus integer H. Milne Edwards, 1837 fide FOREST & GUINOT (1961: 162). — SEURAT, 1934: 58 (NOBILI's
material).
FAMILY GRAPSIDAE
SUBFAMILY GRAPSINAE
Geograpsus crinipes (Dana, 1851)
DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti); Tuamotu (Makatea, Makemo, Mataiva, Moruroa,
Pukapuka, Raroia, Taiaro, Takapoto).
REFERENCES. — Geograpsus crinipes - HELLER, 1865: 48 (Tahiti). — NOBILI, 1907: 404 (Pukapuka). — RATHBUN,
1907: 28 (Makemo). — SENDLER, 1923: 32 (Makatea). — SEURAT, 1934: 52 (Pukapuka). — HOLTHUIS, 1953: 29
(Raroia). — MORRISON, 1954: 9 (Raroia). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). —
MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). —
GUINOT, 1985: 453 (List). — SALVAT & RICHARD, 1985: 359, 360 (Takapoto). — POUPIN, 1994a: 57, fig. 53, pl. 6h
(Moruroa, Nuku Hiva, Taiaro, Takapoto).
Geograpsus grayi (H. Milne Edwards, 1853)
DISTRIBUTION. — Society (Tahiti); Tuamotu (Makatea, Nukutipipi, Raroia).
REFERENCES. — Geograpsus grayi - KINGSLEY, 1880c: 196 (Tahiti). — ORTMANN, 1894: 707 (Tahiti). — SENDLER,
1923: 32, pl. 21, fig. 6 (Makatea). — SEURAT, 1934: 52 (Tahiti). — HOLTHUIS, 1953: 30 (Raroia). — MORRISON, 1954:
11 (Raroia). — BANERDJEE, 1960: 159 (Tahiti). — FoREsT & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). —
MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea). — GUINOT, 1985: 453 (List). — MERSCHARDT-SALVAT, 1991: 40
(Nukutipipi). — SALVAT F. & B., 1992: 5 (Nukutipipi).
Geograpsus stormi De Man, 1895
DISTRIBUTION. — Marquesas (Nuku Hiva).
REFERENCES. — Geograpsus lividus stormi de Man - RATHBUN, 1907: 29 (Nuku Hiva). — Geograpsus stormi -
BANERDIJEE, 1960: 167 (Syn.). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu, Marquesas"). —
GUINOT, 1985: 453 (List). — POUPIN, 1994a: 58, fig. 54, pl. 7a (Nuku Hiva).
EE eee
67
Grapsus depressus Heller, 1862
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Grapsus depressus Heller 1862: 521 (Tahiti; to our knowlege this species has never been mentioned
after its description).
Grapsus longitarsis Dana, 1851
DISTRIBUTION. — Society; Tuamotu (Fakarava, Hao, Hikueru, Makatea, Mataiva, Rangiroa, Raroia, Taiaro, Takapoto,
Tike).
REFERENCES. — Grapsus longitarsis Dana, 1851: 249; 1852b: 339; 1855, pl. 21, fig. 4a-d (Tuamotu). — RATHBUN,
1907: 28 (Fakarava, Rangiroa, Tikei). — HOLTHUIS, 1953: 31 (Raroia). — MORRISON, 1954: 7 (Raroia). — BANERJEE,
1960: 144, fig. 1b, 2h-n (Society, Raroia). — FOREST & GUINOT, 1961: 152, fig. 160a-b, 161, pl. 18, fig. 2 (Hikueru);
1962: 72 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva,
Takapoto). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 59, fig. 55, pl. 7b
(Hao, Hikueru, Taiaro). — Grapsus strigosus - NOBILI, 1907: 404 (Hao) not G. strigosus (Herbst, 1799) synonym of
Grapsus albolineatus Lamarck, 1818 in BANERJEE (1960: 147) = G. longitarsis fide FOREST & GUINOT (1961: 152).
Grapsus tenuicrustatus (Herbst, 1783)
DISTRIBUTION. — Gambier (Kamaka); Marquesas (Nuku Hiva); Society (Moorea, Tahiti); Tuamotu (Fakarava, Hao,
Hikueru, Makatea, Makemo, Mataiva, Moruroa, Rangiroa, Raroia, Taiaro, Takapoto).
REFERENCES. — Grapsus pictus - DANA, 1852b: 336; 1855, pl. 21, fig.1 (Tuamotu). — Grapsus grapsus - NOBILI,
1907: 404 (Hao, Kamaka) not Grapsus grapsus Linné, 1758 = G. tenuicrustatus fide FOREST & GUINOT (1961: 154). —
Grapsus grapsus tenuicrustatus - RATHBUN, 1907: 27 (Fakarava, Makemo, Rangiroa). — Grapsus gracilipes -
SENDLER, 1923: 31(Makatea). — Grapsus gracillimus Sendler, 1923: 32, pl. 21, fig. 5 (Makatea). — Grapsus
tenuicrustatus - HOLTHUIS, 1953: 31 (Raroia). — CHABOUIS L. & F., 1954: 91, fig. 6 (French Polynesia). —
Morrison, 1954: 9 (Raroia). — BANERJEE, 1960: 134, fig. 1a, 2a, c-g (Raroia; Syn.). — FOREST & GUINOT, 1961: 154
(Hikueru; Syn.); 1962: 72 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea,
Mataiva, Moorea, Tahiti, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 453 (List). — SALVAT &
RICHARD, 1985: 360 (Takapoto). — PARDON, 1992: 82, photograph (Tuamotu). — POUPIN, 1994a: 60, fig. 56, pl. 7c
(Nuku Hiva, Tahiti, Taiaro, Takapoto). — The following works refer also probably to this species: Grapsus maculatus
- KINGSLEY, 1880c: 192 (Tahiti) = G. grapsus fide ORTMANN (1894: 703). — Grapsus grapsus - SENDLER, 1923: 30
(Makatea). —BOONE, 1934: 178, pl. 90 ("Anaho Bay” = Nuku Hiva). — CHEVALIER et al., 1968: 95, 138 (Moruroa). —
BAGNIS & CHRISTIAN, 1983: 112-113, photograph (Tuamotu). —Grapsus albolineatus - BONVALLOT et al., 1994: 137,
photograph, (Tuamotu; det. according to the photograph) - SYNONYMS - Grapsus pictus Latreille, 1802-1803; Grapsus
gracilipes H. Milne Edwards, 1853; Grapsus gracillimus Sendler, 1923.
REMARK. — BANERDIEE (1960: 139) mentions that Grapsus grapsus (Linné, 1758) is solely Atlantic. More recently,
MANNING & HOLTHUIS (1981: 233) have also mentioned it from the Eastern Pacific.
Leptograpsus variegatus (Fabricius, 1793)
DISTRIBUTION. — Austral (Rapa); Marquesas.
REFERENCES. — Leptograpsus variegatus - DE MAN, 1890: 84 (Marquesas; with regard of L. ansoni H. Milne Edwards,
1853). — GRIFFIN, 1973: 461, fig. 1-6 (Syn.) - SYNONYMS - Leptograpsus ansoni H. Milne Edwards, 1853 - NEW
MATERIAL - March 1995, Coll. and det. J. POUPIN (Rapa, Haurei bay, very common).
Metopograpsus messor (Forskal, 1775)
DISTRIBUTION. — Society (Bora Bora, Tahiti).
REFERENCES. — Metopograpsus messor - KINGSLEY, 1880c: 190 (Tahiti). — MIERS, 1886: 258 (Tahiti). — RATHBUN,
1907: 29 (Bora Bora). — PESTA, 1913: 61 (Tahiti). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 61 (Text).
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Metopograpsus thukuhar (Owen, 1839)
DISTRIBUTION. — Austral (Tubuai); Gambier (Mangareva); Society (Moorea, Tahiti); Tuamotu (Hao, Taiaro).
REFERENCES. — Metopograpsus thukuar - STIMPSON, 1858b: 101; 1907: 114 [47] (Tahiti). — HELLER, 1865: 43
(Tahiti). — Nosmul, 1907: 404 (Hao, "Rikitea" = Mangareva). — SEURAT, 1934: 59 (Mangareva). — BANERJEE, 1960:
186, fig. 6f-g (Tahiti). — FOREST & GUINOT, 1961: 155, fig. 162, 167 (Tahiti); 1962: 72 (Biogeography "Tahiti-
Tuamotu"). — GUINOT, 1985: 453 (List). — MARQUET, 1988: 90, fig. 48, tab. 23; 1991: 130, tab. 1-2; 1993: tab. 1, 3
(Mangareva, Moorea, Tahiti, Tubuai). — POUPIN, 1994a: 61, fig. 57, pl. 7d (Hao, Mangareva, Tahiti, Taiaro).
Pachygrapsus fakaravensis Rathbun, 1907
DISTRIBUTION. — Tuamotu (Fakarava, Makatea, Mataiva, Taiaro, Takapoto).
REFERENCES. — Pachygrapsus fakaravensis Rathbun, 1907: 29, pl. 5, fig. 1, pl. 9, fig. 6, 6a (Fakarava). — FOREST &
GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva,
Takapoto). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 62, fig. 58, pl. 7e (Taiaro).
Pachygrapsus minutus A. Milne Edwards, 1873
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru, Makatea, Mataiva, Takapoto, Tikehau).
REFERENCES. — Pachygrapsus minutus - FOREST & GUINOT, 1961: 155 (Hikueru, Tahiti); 1962: 72 (Biogeography
"Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 25; 1977b: 213; 1985: 462 (Moorea); 1989:
113 (Tikehau). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9
(Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 453 (List) - RELEVANT MATERIAL - Pachygrapsus
aff. minutus - NAIM, 1980a: 55, annex 1, tab. 3 (Moorea; very small adult specimens, possibly of a new species). —
GUINOT, 1985: 453 (List).
Pachygrapsus planifrons De Man, 1888
DISTRIBUTION. — Tuamotu (Fakarava, Raroia).
REFERENCES. — Pachygrapsus planifrons - HOLTHUIS, 1953: 31 (Raroia). — MORRISON, 1954: 7 (Raroia). — FOREST
& GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). — Pachygrapsus longipes -
RATHBUN, 1907: 30 (Fakarava) - SYNONYMS - Pachygrapsus longipes Rathbun, 1893 (in TESCH, 1918, p. 78; with
uncertainty).
Pachygrapsus plicatus (H. Milne Edwards, 1837)
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava, Hikueru, Makatea, Makemo, Mataiva, Raroia, Taiaro,
Takapoto).
REFERENCES. — Pachygrapsus plicatus - KINGLEY, 1880c: 200 (Tahiti). — RATHBUN, 1907: 29 (Fakarava, Makemo).
— HOLTHUIS, 1953: 32 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST & GUINOT, 1961: 154 (Hikueru); 1962:
72 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti,
Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 453 (List). — SALVAT &
RICHARD, 1985: 359 (Takapoto). — POUPIN, 1994a: 63, fig. 59, pl 7f (Hikueru, Taiaro).
Planes cyaneus Dana, 1851
DISTRIBUTION. — Austral (Neilson bank).
REFERENCES. — Planes cyaneus - NEW MATERIAL - April 1995, coll. J. POUPIN, det. A. CROSNIER (Neilson bank; on a
drifting buoy with cirripeds).
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SUBFAMILY VARUNINAE
Hemigrapsus crenulatus (H. Milne Edwards, 1837)
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Heterograpsus crenulatus Guérin - NOBILI, 1907: 405 (Tahiti; in Paris, NOBILI's material is well
recorded under Hemigrapsus crenulatus MNHN B12830). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-
Tuamotu"). — Hemigrapsus crenulatus (H. Milne Edwards) - BENNETT, 1964: 81 (Syn.). — GUINOT, 1985: 453 (List).
Pseudograpsus albus Stimpson, 1858
DISTRIBUTION. — Tuamotu (Fakarava, Raroia).
REFERENCES. — Pseudograpsus albus - RATHBUN, 1907: 32 (Fakarava). — HOLTHUIS, 1953: 32 (Raroia). —
Morrison, 1954: 10 (Raroia). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). —GUuINOT, 1985:
453 (List).
Ptychognathus crassimanus Finnegan, 1931
DISTRIBUTION. — Marquesas - Freshwater.
REFERENCES. — Ptychognathus crassimanus Finnegan, 1931: 649 (Marquesas).
REMARK. — It seems that this species was never recorded after its description. In particular, it does not appear in the
works by MARQUET (1988, 1991, 1993), who has only collected Ptychognathus easteranus (det. HOLHTUIS), in the
Marquesas rivers.
Ptychognathus easteranus Rathbun, 1907
DISTRIBUTION. — Austral (Rurutu); Marquesas (Hiva Oa) - Freshwater.
REFERENCES. — Ptychognathus easteranus - MARQUET, 1988: 90, fig. 48, tab. 23; 1991: 132, tab. 1-2; 1993: tab. 1, 3
(Hiva Oa, Rurutu).
Ptychognathus intermedius (de Man, 1879)
DISTRIBUTION. — Society (Tahiti) - Freshwater.
REFERENCES. — Ptychognathus intermedius - ORTMANN, 1894: 711 (Tahiti). — FOREST & GUINOT, 1962: 72
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List).
Thalassograpsus harpax (Hilgendorf, 1892)
DISTRIBUTION. — French Polynesia.
REFERENCES. — Thalassograpsus harpax - GUINOT, 1985: 453 (List only; origin not found).
Varuna litterata (Fabricius, 1798)
DISTRIBUTION. — Society (Moorea, Tahiti) - Fresh & Brackish water.
REFERENCES. — Varuna litterata - MARQUET, 1988: 90, fig. 48, tab. 3; 1991: 133, tab. 1-2; 1993: tab. 1, 3 (Moorea,
Tahiti). — POUPIN 1994a: 67, fig. 63, pl. 8b (Tahiti).
SUBFAMILY SESARMINAE
Chasmagnathus subquadratus Dana, 1851
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Chasmagnathus subquadratus - ORTMANN, 1894: 728 (Tahiti). — ForREsT & GUINOT, 1962: 72
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List).
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REMARK. — Species described by par DANA (1851: 251) from an uncertain locality "Novi-Zealandiae ? Novi-
Hollandiae orientalis 2", not very often cited.
Cyclograpsus integer H. Milne Edwards, 1837
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava, Hikueru, Kaukura?, Mataiva, Raroia, Taiaro,
Takapoto).
REFERENCES. — Epigrapsus politus - NOBILI, 1907: 407 (Hikueru, Kaukura) not Epigrapus politus Heller, 1862 = C.
integer fide FOREST & GUINOT (1961: 160). — Cyclograpsus parvulus - RATHBUN, 1907: 36 (Fakarava). — HOLTHUIS,
1953: 32 (Raroia). — MORRISON, 1954: 5 (Raroia). — Cyclograpsus integer - FOREST & GUINOT, 1961: 160, fig. 175a-
c (Hikueru; Syn.); 1962: 74 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Mataiva,
Moorea, Tahiti, Takapoto). — GUINOT, 1985: 454 (List). — POUPIN, 1994a: 56, fig. 52, pl. 6f (Hikueru, Taiaro) -
SYNONYMS - Cyclograpsus parvulus de Man, 1897.
Cyclograpsus longipes Stimpson, 1858
DISTRIBUTION. — Society (Tahiti); Tuamotu (Makemo, Marutea South, Raroia).
REFERENCES. — Cyclograpsus longipes - RATHBUN, 1907: 36 (Makemo, Tahiti). — HOLTHUIS, 1953: 32 (Raroia). —
Morrison, 1954: 5 (Raroia). — FOREST & GUINOT, 1961: 160 (Marutea South); 1962: 72 (Biogeography "Tahiti-
Tuamotu"). — GUINOT, 1985: 454 (List).
Labuanium trapezoideum (H. Milne Edwards, 1837)
DISTRIBUTION. — Society (Moorea, Raiatea, Tahiti) - Freshwater.
REFERENCES. — Sesarma trapezoidea - GUERIN-MENEVILLE, 1838: 14 (Tahiti). — SEURAT, 1934: 51 (Tahiti). —
FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). — MARQUET, 1988: 90, fig. 48, tab. 23; 1991: 133, tab.
1-2; 1993: tab. 1, 3 (Moorea, Tahiti). — Sesarma (Sesarma) trapezoidea - NOBILI, 1907: 405 (Tahiti). — EDMONSON,
1951: 237, fig. 33b (Raiatea, Tahiti). — FOREST & GUINOT, 1961: 157, fig. 164a-b, 165 (Tahiti). — Sesarma (Sesarma)
trapezoideum - RATHBUN, 1907: 33 (Tahiti). — Labuanium trapezoideum - SERENE & SOH, 1970: 402, 406 (Syn.). —
GUINOT, 1985: 454 (List).
REMARK. — Labuanium rotundatum (Hess, 1865) is also recorded in Polynesia by SAKAI (1976: 663; distribution only
"Micronesia, Polynesia"; cited afterwards by GUINOT, 1985: 454). SAKAI must consider the Polynesia s./., with about 10
states, including French Polynesia. It is doubtful that this species have been really collected in French Polynesia since
we have not find any mention of it in TESCH (1917: 193), who gives a detailed distribution, or in SERENE & SOH (1970:
402, 406), when they have established the genus Labuanium. For the moment, it thus seems better to exclude it from the
area.
Metasesarma rousseauxi granularis Heller, 1862
DISTRIBUTION. — Gambier (Tarauru-Roa); Society (Tahiti ); Tuamotu (Hikueru).
REFERENCES. — Metasesarma granularis Heller, 1862: 522 (Tahiti). —Metasesarma rugulosa Heller, 1865: 65
(Tahiti; cf. Remark). — Metasesarma rousseauxi H. Milne Edwards - ? ORTMANN, 1894: 717 (Tahiti). — ? HOLTHUIS,
1953: 33 ("Taravao" = Tahiti). — Sesarma (Metasesarma) rousseauxi - NOBILI, 1907: 405 (Tarauru-Roa). —
Metasesarma rousseauxi granularis - FOREST & GUINOT, 1961: 158, fig. 168, 169, 174a-b (Hikueru, “Papenoo"” =
Tahiti; Syn.); 1962: 72 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 454 (List).
REMARK. — FOREST & GUINOT (1961: 158) writes "M. rousseauxi granularis, décrit en 1862 de Tahiti par HELLER, qui,
en 1865, substituait 4 ce nom, sans raison apparente, celui de M. granulosa, a été mis en synonymie avec Metasesarma
rousseauxi H. Milne Edwards, 1853, par DE MAN (1889, p. 439)". The same authors recognise differences between H.
MILNE EDWARDS' species and the specimens from Tahiti and Tuamotu, which they attribute to the subspecies granularis
Heller. According to the location, ORTMANN's and HOLTHUIS' references should be also attributed to this subspecies.
Sarmatium crassum Dana, 1851
DISTRIBUTION. — Society (Tahiti).
aS. Saeco eer
al
REFERENCES. — Sarmatium crassum - GUINOT, 1985: 454 (List; origin ?). — DAVIE, 1992: 81, fig. 1a, 2, 3a-c (Tahiti).
Sesarma angustifrons A. Milne Edwards, 1869
DISTRIBUTION. — Society (Moorea, Tahiti).
REFERENCES. — Sesarma (Sesarma) angustifrons - DEMAN, 1889: 432, pl. 10, fig. 10 (Tahiti). — NoBILI, 1907: 405
(Tahiti). — SEURAT, 1934: 51 (Tahiti). — Sesarma angustifrons - FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-
Tuamotu"). — GUINOT, 1985: 454 (List). — MARQUET, 1988: 90, fig. 48, tab. 23; 1991: 133, tab. 1-2; 1993: tab. 1, 3
(Moorea, Tahiti).
Sesarma jacquinoti Ortmann, 1894
DISTRIBUTION. — Society (Tahiti).
REFERENCES. — Sesarma jacquinoti Ortmann, 1894: 718 (Tahiti). — FOREST & GUINOT, 1962: 72 (Biogeography
"Tahiti-Tuamotu”). — GUINOT, 1985: 454 (List).
SUBFAMILY PLAGUSIINAE
Percnon abbreviatum (Dana, 1851)
DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti); Tuamotu (Raroia).
REFERENCES. — Acanthopus abbreviatus Dana, 1851: 252; 1852b: 373; 1855, pl. 23, fig. 11a-c (Tahiti). — Percnon
affinis - NoBILI, 1907: 406 (Mangareva), pro parte not P. affine H. Milne Edwards, 1853 = P. abbreviatum fide FOREST
& GUINOT (1961: 164). — Percnon abbreviatum - HOLTHUIS, 1953: 33 (Raroia). — MORRISON, 1954: 16 (Raroia). —
EDMONSON, 1959: 195, fig. 25c, 26a-c (Syn.). — FOREST & GUINOT, 1961: 164 (Mangareva; Syn.); 1962: 74
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (Raroia); 1985: 454 (List).
Percnon affine (H. Milne Edwards, 1853)
DISTRIBUTION. — Marquesas; Gambier (Mangareva); Society (Tahiti); Tuamotu (Makatea, Mataiva).
REFERENCES. — Percnon affinis - NoBILI, 1907: 406 ("Gatavake” = Mangareva), pro parte some sp. are P.
abbreviatum — FOREST & GUINOT, 1961: 164 (""Gatavake” = Mangareva; Syn.); 1962: 74 (Biogeography "Tahiti-
Tuamotu"). — Percnon pilimanus - BOONE, 1934: 181, pl. 92-94 (Tahiti). — FOREST & GUINOT, 1962: 74
(Biogeography "Tahiti-Tuamotu, Marquesas”). — Percnon affine - CROSNIER, 1965: 88 (Tuamotu; Syn.). —
MONTEFORTE, 1984: 172, annex 1, tab. a, photograph p. 138a-bas (Makatea, Mataiva, Tahiti). — DELESALLE, 1985: 289
(Mataiva). — GUINOT, 1985: 454 (List) - SYNONYMS - Percnon pilimanus (A. Milne Edwards, 1873).
Percnon guinotae Crosnier, 1965
DISTRIBUTION. — .Marquesas (Hiva Oa).
REFERENCES. — Percnon guinotae - NEW MATERIAL - February 1996, Coll. & det. J. POUPIN (Hiva Oa).
REMARK. — The presence of this species in French Polynesia was assumed in POUPIN (1994a: 64). It is confirmed here
with a specimen collected in the Marquesas.
Percnon planissimum (Herbst, 1804)
DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Tahiti); Tuamotu (Fakarava, Hao, Makatea,
Marutea South, Mataiva, Moruroa, Raraka, Taiaro, Tikehau).
REFERENCES. — Acanthopus planissimus - DANA, 1852b: 372 (Raraka, Tahiti). — HELLER, 1865: 51(Tahiti). —
Acanthopus tenuifrons H. Milne Edwards, 1853: 180 (Nuku Hiva). — Percnon planissimus - NOBII, 1907: 406 (Hao,
Mangareva, Marutea). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Tahiti). — PEYROT-CLAUSADE,
1989: 113 (Tikehau). — Percnon planissimum - RATHBUN, 1907: 37 (Fakarava). — PESTA, 1913: 64 (Tahiti). —
EDMONDSON, 1959: 197, fig. 25c, 27a-c (Marquesas). — FOREST & GUINOT, 1961: 163 (Marutea South, Tahiti; Syn.);
72
1962: 74 (Biogeography "Tahiti-Tuamotu, Marquesas"). — CHEVALIER et al., 1968: 95, 138 (Moruroa). — GUINOT,
1985: 454 (List). — POUPIN, 1994a: 64, fig. 60, pl. 8a (Mangareva, Taiaro).
Plagusia speciosa Dana, 1851
DISTRIBUTION. — Marquesas; Society (Tahiti); Tuamotu (Ahe and/or Manihi, Hao, Hikueru, Makatea, Makemo,
Mataiva, Raroia, Taiaro, Takapoto).
REFERENCES. — Plagusia speciosa Dana, 1851: 252; 1852b: 369; 1865, pl. 23, fig. 9 ("Waterland" = Ahe and/or
Manihi). — KINGSLEY, 1880c: 223 (Tahiti). — DE MAN, 1890: 89 (Tuamotu). — ORTMANN, 1894: 731 (Tuamotu). —
NoBILI, 1907: 406 (Hao). — RATHBUN, 1907: 36 (Makemo). — SENDLER, 1923: 35 (Makatea). — BOONE, 1934: 185,
pl. 95-96 (Tahiti). — HOLTHUIS, 1953: 34 (Raroia). — MORRISON, 1954: 16 (Raroia). — FOREST & GUINOT, 1961: 162,
fig. 177a-c, 178 (Hao, Hikueru); 1962: 74 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1966a: 48
(Raroia); 1985: 454 (List). — MONTEFORTE, 1984: 172, annex 1, tab. a, photograph p. 138a-haut (Makatea, Mataiva,
Takapoto). — DELESALLE, 1985: 289 (Mataiva). — SALVAT & RICHARD, 1985: 362 (Takapoto). — POUPIN, 1994a: 65,
fig. 61, pl. 7g (Hao, Hikueru, Taiaro).
REMARK. — Plagusia immaculata Lamarck, 1818 is erroneously cited from Tahiti and the Tuamotu by DAI & YANG
(1991: 563). This error must come from a quick reading of EDMONSON (1959: 194), where "Tuamotus (type locality),
Tahiti” is mentioned under P. immaculata, but for remarks concerning only P. speciosa Dana.
Plagusia tuberculata Lamarck, 1818
DISTRIBUTION. — Austral (Raevavae); Gambier (Kamaka, Mangareva); Marquesas (Nuku Hiva); Tuamotu (Makatea?).
REFERENCES. — Plagusia squamosa - NOBILI, 1907: 406 (Kamaka, Mangareva; inferred only: reference not retrieved in
recent works, and material not found in Paris). — Plagusia depressa tuberculata - FOREST & GUINOT, 1962: 74
(Biogeography "Tahiti-Tuamotu"). — Plagusia depressa - ? MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea) not P.
depressa (Fabricius, 1775) = P. tuberculata cf. Remark. — Plagusia tuberculata - GUINOT, 1985: 454 (List). —
POUPIN, 1994a: 66, fig. 62, pl. 7h (Nuku Hiva, Raevavae, Tuamotu) - SYNONYMS - Plagusia squamosa (Herbst, 1882)
(cf. SAKAI, 1976: 676, then ALCOCK, 1900: 437).
REMARK. — Plagusia depressa (Fabricius, 1775) is a species of the tropical Atlantic (cf. DAWSON, 1987: 42) and cannot
be MONTEFORTE's (1984) material which is more likely P. tuberculata (One specimen of this species is actually
deposited in the CRIOBE collections, Moorea, Coll. and det. MONTEFORTE).
FAMILY PINNOTHERIDAE
Pinnotherelia laevigata A. Milne Edwards & Lucas, 1843
DISTRIBUTION. — Marquesas (Nuku Hiva).
REFERENCES. — Pinnotheralia laevigata - RATHBUN, 1918: 181, fig. 115, pl. 39, fig. 1-3, pl. 40, fig. 1-2 (Marquesas
"Tawhoe" = Taiohae, Nuku Hiva). — SCHMITT et al., 1973: 125 (catalogue "Marquesas Islands").
FAMILY OCYPODIDAE
Macrophthalmus consobrinus Nobili, 1906
DISTRIBUTION. — Gambier (Mangareva) - Brackish water.
REFERENCES. — Macrophthalmus consobrinus Nobili, 1906a: 265; 1907: 408 (Mangareva). — FOREST & GUINOT,
1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). — Macrophthalmus parvimanus - BARNES,
1977: 273, pro parte, only NOBILI's consobrinus, not M. parvimanus Guérin-Méneville, 1834 (cf. Remark).
REMARK. — According to BARNES (1977: 273) NoBILI's Macrophthalmus consobrinus is the same as Macrophthalmus
parvimanus Guérin-Méneville, 1834. However, to check that point, a large sample of M. consobrinus has been recently
73
collected in the Gambier Islands, and it appears that NOBILI's species is valid, and closely related to M. convexus
Stimpson, 1858 (POUPIN, in study).
Macrophthalmus convexus Stimpson, 1858
DISTRIBUTION. — Society (Bora Bora, Tahiti) - Brackish water.
REFERENCES. — Macrophthalmus convexus - ORTMANN, 1894: 745 (Tahiti). — FOREST & GUINOT, 1962: 70
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List) - NEW MATERIAL - Coll. and det. J. POUPIN (Bora Bora).
Macrophthalmus serenei (Seréne, 1983)
DISTRIBUTION. — Tuamotu (Tikehau).
REFERENCES. — Macrophthalmus serenei - NEW MATERIAL - Coll. C. HILY, det. J. POUPIN (Tikehau).
Ocypode ceratophthalma (Pallas, 1772)
DISTRIBUTION. — Gambier? (Mangareva); Society (Scilly, Tahiti); Tuamotu (Fakarava, Makatea, Makemo, Mataiva,
Marutea South?, Rangiroa, Raroia, Taiaro, Takapoto, Tikehau).
REFERENCES. — Ocypode urvillei Guérin-Méneville, 1829: pl. 1, fig. 1, la-b; 1838: 9 (Tahiti). — OWEN, 1839: 80
("Low Islands" = Tuamotu). — NoBILI, 1907: 407 pro parte (Marutea?, cf. Remark sous O. pallidula). — FOREST &
GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453, with a ? (List). — Ocypode
ceratophthalma - STIMPSON, 1858b: 100 [46]; 1907: 108, pl. 12, fig. 2 (Tahiti). — ORTMANN, 1897a: 364 (Syn.). —
RATHBUN, 1907: 26 (Fakarava, Makemo, Rangiroa). — SENDLER, 1923: 21 (Tahiti). — HOLTHUIs, 1953: 28 (Raroia).
— Morrison, 1954: 9 (Raroia). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — SAKAI &
TURKAY, 1976: 86, fig. 13 (Syn.). — MONTEFORTE, 1984: 173, annex 1, tab. a (Makatea, Mataiva, Takapoto). —
DELESALLE, 1985: 289, 303 (Mataiva). — GUINOT, 1985: 453 (List). — HARMELIN-VIVIEN, 1985: 239 (Tikehau). —
SALVAT & RICHARD, 1985: 359 (Takapoto). — POUPIN, 1994a: 68, fig. 64, pl. 8c-d (Tahiti, Taiaro, Scilly). — Ocypode
cordimana - KINGSLEY, 1880b: 186 (Tahiti) not O. cordimana Desmaret, 1825 = O. urvillei, synonym of O.
ceratophthalma, fide ORTMANN (1897a: 366). — ? Oxypode (sic) - CHARLEUX, 1986: 81, photograph (French
Polynesia), det. according to the photograph.
Ocypode cordimana Desmaret, 1825
DISTRIBUTION. — Society (Bora Bora, Tahiti).
REFERENCES. — Ocypode cordimana - BOONE, 1934: 191, pl. 99-100 (Bora Bora). — FOREST & GUINOT, 1962: 70
(Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 69, fig. 65, pl. 8e (Tahiti). — Not
Ocypode cordimana - KINGSLEY, 1880b: 186 (Tahiti) fide ORTMANN (1897a: 366) = O. urvillei, synonym of O.
cerathophthalma.
REMARK. — BOONE mentions this species in the Tuamotu ("Paumotus") after RATHBUN (1907: 26). In fact it is O.
cerathophthalma that RATHBUN has cited from this archipelago.
Ocypode pallidula Jacquinot 1852
DISTRIBUTION. — Gambier (Aukena, Mangareva); Tuamotu (Marutea South?, Moruroa).
REFERENCES. — Ocypode pallidula Jacquinot, 1852, pl. 6, fig. 1a (Mangareva). — SAKAI & TURKAY, 1976: 87, fig. 14-
15 ("Rikitea" = Mangareva; type material, Syn.). — JONES, 1988: 34 (Syn.). — POUPIN, 1994a: 70, fig. 66, pl. 8f
(Aukena). — Ocypode cordimana (Junior) - JACQUINOT & LUCAS, 1853: 64 (Mangareva) not O. cordimana Desmaret,
1825 = O. pallidula fide SAKAI & TURKAY (1976: 87). — Ocypode urvillei - NOBILI, 1907: 407, pro parte (“Rikitea" =
Mangareva, Marutea?). — SEURAT, 1934: 52 (Mangareva; NOBILI's material) - The two references, not O. urvillei
Guérin-Méneville, 1829, synonym of O. cerathophthalma = O. pallidula (cf. Remark). — Ocypode laevis - CHEVALIER
et al., 1968: 109 (Moruroa) - NEW MATERIAL - Coll. B. SALVAT, 1966, det. J. POUPIN (Moruroa) - SYNONYMS - Ocypode
laevis Dana, 1852.
REMARK. — After ORTMANN (1897a: 366), Ocypode pallidula, was usually considered as the same as O. urvillei (= O.
cerathophthalma). Its validity was re-established by SAKAI & TURKAY (1976). The material mentioned in NOBILI (1907)
74
and SEURAT (1934), under O. urvillei has been collected at Marutea South and Mangareva. The specimens from this
second island are in fact O. pallidula (verification in the collections of Paris: 1 sp. labelled "Ocypoda urvillei Guér.,
Seurat coll. 1905, Bouvier dét., G. Nobili vérif. 1906", MNHN B11841, is a real O. pallidula). The specimens from
Marutea South could reasonably be O. cerathophthalma, very common in the Tuamotu.
Uca chlorophthalmus crassipes (Adams & White, 1848)
DISTRIBUTION. — Austral (Raevavae); Marquesas; Society (Bora Bora, Maupiti, Raiatea, Tahiti) - Brackish water.
REFERENCES. — Gelasimus latreillei H. Milne Edwards, 1852: 114, pl. 4, fig. 20, 20a (Bora Bora). — Gelasimus
pulchellus Stimpson, 1858b: 100 [46]; 1907: 107, pl. 15, fig. 1 (Tahiti). — Gelasimus gaimardi - HELLER, 1865: 38
(Tahiti). — Uca chlorophthalmus - NoBILI, 1907: 408 (""Taravao" = Tahiti). — Uca gaimardi - RATHBUN, 1907: 26
(Bora Bora, Tahiti). — HOLTHUIS, 1953: 29 ("Taravao" = Tahiti). — CRANE, 1957: 74, 78 (Bora Bora, Raiatea, Tahiti).
— FOREST & GUINOT, 1961: 140, fig. 140-145, 153, 156a-b (Tahiti); 1962: 70 (Biogeography "Tahiti-Tuamotu"). —
Gelasimus (Uca) chlorophthalmus - SEURAT, 1934: 60 (Tahiti). — Uca (Amphiuca) chlorophthalmus crassipes -
CRANE, 1975: 98, 102, 599, fig. 13-14, 26c, 31c, 37h, 39a-b, 56c, 60 I-m, 68a-b, 81g, 83a, 99, pl. 15 a-f, 46b (Bora
Bora, Raiatea, Tahiti, Marquesas p. 599; Syn.). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 71, fig. 67, pl. 8g
(Maupiti, Raevavae, Tahiti) - SYNONYMS - Uca gaimardi H. Milne Edwards, 1852.
Uca tetragonon (Herbst, 1790)
DISTRIBUTION. — Austral (Rapa); Gambier (Mangareva); Society (Bora Bora, Raiatea, Tahiti); Tuamotu (Moruroa,
Napuka, Raroia).
REFERENCES. — Gelasimus duperreyi Guérin-Méneville, 1829, pl. 1, fig. 2, 2a (Bora Bora). — Gelasimus tetragonon -
GUERIN-MENEVILLE, 1838, pl. 1, fig. 2, 2a (Bora Bora) pro parte. — HELLER, 1865: 37 (Tahiti). — KINGSLEY, 1880a:
143, pl. 9, fig. 11 (Tahiti). — MIERS, 1886: 243 (Tahiti). — DE MAN, 1891: 24, pl. 2, fig. 6 (Tahiti). — ORTMANN,
1894: 754 (Tahiti). — Uca tetragonon - NoBILI, 1907: 408 ("Rikitea, Gatavake" = Mangareva). — RATHBUN, 1907: 26
(Bora Bora). — HOLTHUIs, 1953: 29 (Raroia). — MORRISON, 1954: 8 (Raroia). — CRANE, 1957: 79 (Bora Bora). —
Gelasimus (Uca) tetragonon - SEURAT, 1934: 59 (Mangareva). — Uca tetragonum - FOREST & GUINOT, 1962: 70
(Biogeography "Tahiti-Tuamotu”). — Uca (Thalassuca) tetragonon - CRANE, 1975: 77, 81, 596, fig. 37d, 63a-b, 81f,
82e, 99, pl. 13 (Bora Bora, Raiatea, Raroia, Tahiti; Syn.). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 72, fig. 68,
pl. 8h (Mangareva, Moruroa, Napuka, Rapa).
REMARK. — FOREST & GUINOT (1962: 70) mention U. dussumieri (H. Milne Edwards) from French Polynesia, probably
after ORTMANN (1894; Gelasimus dussumieri, Tahiti, p. 755). ORTMANN's reference is cited by CRANE (1975) for two
subspecies: Uca (Deltuca) dussumieri spinata (specimens from Java and Singapore) and Uca (Deltuca) dussumieri
dussumieri (specimens from the Philippines and Mindanao). ORTMANN's Tahitian U. dussumieri do not appear in
CRANE, who clearly indicates (p. 437) that Uca dussumieri does not occur in French Polynesia. ORTMANN's "U.
dussumieri", if they exist, must probably be, either U. chlorophthalmus, or U. tetragonon.
FAMILY CRYPTOCHIRIDAE
Cryptochirus coralliodytes Heller, 1861
DISTRIBUTION. — Tuamotu (Makatea, Marutea South, Marokau).
REFERENCES. — Cryptochirus coralliodytes - NOBILI, 1907: 409 (Marutea South, Marokau). — SENDLER, 1923: 41
(Makatea). — FOREST & GUINOT, 1962: 74 (Biogeography "Tahiti-Tuamotu"). — SEURAT, 1934: 60 (Marokau,
Marutea South). — GUINOT, 1985: 454 (List). — KRopp, 1988: 873 (Revision of this species but without mention of
French Polynesia).
Hapalocarcinus marsupialis Simpson, 1859
DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Takapoto).
75
REFERENCES. — Hapalocarcinus marsupialis - KRopp & BIRKELAND, 1981: 629, tab. 5 (Moorea, Takapoto). —
ODINETZ, 1983: 29, 205 (Moorea, Tahiti, Takapoto). — GUINOT, 1985: 454 (List). — ODINETZ-COLLART & RICHER DE
ForRGES, 1985: 201 (Moorea and/or Tahiti).
FAMILY HYMENOSOMATIDAE
Elamena mathaei (Desmaret,1825)
DISTRIBUTION. — Tuamotu (Tikehau).
REFERENCES. — Elamena mathaei - PEYROT-CLAUSADE, 1989: 113 (Tikehau).
FAMILY INCERTAE SEDIS
Daira perlata (Herbst, 1790)
DISTRIBUTION. — Society (Moorea, Raiatea, Tahiti); Tuamotu (Makatea, Mataiva, Hao, Hikueru, Raroia, Taiaro,
Takapoto, Tikehau).
REFERENCES. — Daira perlata - NOBILI, 1907: 392 (Hao). — RATHBUN, 1907: 44 (Tahiti). — SENDLER, 1923: 38
(Makatea). — BOOng, 1934: 129, pl. 69 (Raiatea, Tahiti). — HOLTHUIS, 1953: 19 (Raroia). — CHABOUIS L. & F., 1954:
91, fig. 4 (French Polynesia). — MORRISON, 1954: 15 (Raroia). — FOREST & GUINOT, 1961: 119 (Hikueru, Tahiti);
1962: 68 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212
(Moorea); 1989: 112, 115 (Moorea, Tikehau). — MONTEFORTE, 1984: 170, annex 1, tab. a, photograph p. 137a
(Makatea, Mataiva, Moorea, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289, 305 (Mataiva). — GUINOT, 1985:
453 (List). — SALVAT & RICHARD, 1985: 362 (Takapoto). — SALVAT, 1986b: 72, photograph (French Polynesia). —
POUPIN, 1994a: 51, fig. 47, pl. 6b (Hikueru, Taiaro).
REMARK. — This species has sometimes been classified in the Zalasiinae Seréne, 1968 (cf. SAKAI, 1976: 513).
Parapleurophrycoides roseus Nobili, 1906
DISTRIBUTION. — Tuamotu (Marutea South).
REFERENCES. — Parapleurophrycoides roseus Nobili, 1906a: 264; 1907: 402, pl. 2, fig. 5 (Marutea; gen. nov. and sp.
nov. described from a very small specimen, 1.3x1.7mm).
REMARK. — For this species, and the following, FOREST & GUINOT (1962: 41) wnite: "...nous les considérons comme
des juvéniles difficilement identifiables."
Platyozius perpusillus Nobili, 1906
DISTRIBUTION. — Tuamotu (Hao).
REFERENCES. — Platyozius perpusillus Nobili, 1906a: 264; 1907: 401 (Hao; described from a very small specimen,
1.45x1.75mm).
REMARK. — In SAKAI (1976: 535), Platyozius Borradaile, 1902 is the same as Eucrate de Haan, 1835 (Goneplacidae).
76
DISCUSSION
NUMBER OF POLYNESIAN SPECIES
A total of 401 littoral or sublittoral species are recorded in this work. The number by infra-order and
family is computed on table 1. The Brachyura clearly prevail, with 78% of the species, followed by the
Anomura (18%), and the Palinura (4%).
Within the crabs, the Xanthidae account for 123 species, distributed in 5 main subfamilies:
Liomerinae, Actaeinae, Zoziminae, Xanthinae, and Chlorodiinae. The Portunidae account for 54 species,
including 6 recorded for the first time from determinations made by MOOSA and CROSNIER: Portunus
macrophthalmus, P. orbitosinus, Thalamita danae, T. macropus, T. mitsiensis, and T. philippinensis; half of
the sublittoral species belong to this family. Two other families are also well represented, the Grapsidae,
with 35 species, and the Trapeziidae, with 20 species. From these two families, Percnon guinotae, Planes
cyaneus and Quadrella maculosa, are recorded for the first time. With respect to the list presented 10 years
ago by GUINOT (1985), excluding the species mentioned erroneously, or not fully determined, about 60
species are added to the Polynesian Brachyura, and approximately a hundred, if the deep species are
included.
Within the Anomura, the Diogenidae account for 40% of the species with three main genera:
Calcinus, Clibanarius, and Dardanus. Calcinus guamensis, C. imperialis, and Dardanus australis, are now
recorded in French Polynesia. The porcellanids account for about 25% of the species, the single genus
Petrolisthes representing 12 species out of 17. The Albuneidae are represented by only one species,
Albunea speciosa, which was previously thought to be endemic from the Hawaiian islands.
Only 14 palinurids are recorded in French Polynesia, of which 2 only by larvae (Palinurellus
wieneckii and Arctides regalis).
SPECIES ERRONEOUSLY RECORDED IN FRENCH POLYNESIA
Fourteen species have been erroneously recorded in French Polynesia. They are: Panulirus ornatus
(cf. under P. versicolor), Panulirus polyphagus (cf. under P. pascuensis), Dynomene sinense (cf. under D.
praedator), Ashtoret granulosa (cf. under A. picta), Lophozozymus incisus (cf. under L. superbus),
Labuanium rotundatum (cf. under L. trapezoideum), Plagusia immaculata (cf. under P. speciosa), Uca
dussumieri (cf. under U. tetragonon), Pachygrapsus transversus, Dotilla fenestrata, Ocypode macrocera,
Ocypode platytarsis, and two freshwater crabs (Potamonidae).
Pachygrapsus transversus Gibbes, 1850 is recorded from Tahiti by KINGSLEY (1880c: 199)
(Tahiti). According to HOLTHUIS & GOTTLIED (1958: 102) this record is obviously false, P. transversus
being an Atlantic species. This conclusion is later supported by MANNING & HOLTHUIS (1981: 235), who
report however the species in the Pacific, but only along the American coasts.
The ocypodid Dotilla fenestrata Hilgendorf, 1869, is listed by GUINOT (1985: 453) after KROPP &
BIRKELAND (1981). It is probably a mistake, because this species is not mentioned in that work, and, to
our knowledge, has never been reported, elsewhere, from French Polynesia. Two other ocypodids, with an
uncertain status, are also erroneously reported from Tahiti, by HELLER (1865: 42): Ocypode macrocera (H.
Milne Edwards, 1837) and Ocypode platytarsis (H. Milne Edwards, 1852) (see ORTMANN, 1897a: 362).
qd
HELLER has mentioned two potamonids crabs in Tahiti: Thelphusa wiillerstorfi, described as a new
species in 1862 (p. 520); and Thelphusa leschenaudii (H. Milne Edwards, 1853) (in HELLER, 1865: 32).
RATHBUN (1904: 287) places these two references under a single species Potamon (Potamon)
hydrodromus (Herbst, 1796) and writes (p. 289): "il est douteux que cette espéce ou quelqu'autre habite
Tahiti". Since the recent works by MARQUET (1988, 1991, 1993), who has intensively sampled the
freshwater Polynesian fauna, it is almost certain that the Potamonidae are not represented in French
Polynesia.
IMPROVEMENT AND CORRECTION OF THIS LIST
This bibliographic compilation is of course tentative and certainly does not account for all the
species living in French Polynesia. When new collections become available, other species will undoubtedly
be added to the present list. Moreover, despite a careful research, it is possible that a few works, recording
additional species, have passed undetected. Right now, several species listed here deserve a particular
attention, either because their presence in French Polynesia remained to be confirmed, or because their
identification, or taxonomic status, are uncertain.
Twelve species of this list could have been erroneously recorded from French Polynesia. They are
known only by larvae (Palinurellus wieneckii, Arctides regalis), are mentioned with doubt, or in an
ambiguous way (Petrolisthes militaris, Porcellana mitra, Porcellana monilifera, Ozius truncatus), appear
only in a part "Distribution", the origin of the French Polynesian material remaining unknown (Neoliomera
insularis, Gaillardiellus rueppelli, Macromedaeus distinguendus, Thalassograpsus harpax), or, are cited
from French Polynesia only because of the large geographic distribution of the species, without real
collections in the field (Schizophrys aspera, Aethra scruposa).
For a score of species the revision of the material would be particularly interesting. They are
Coenobita cavipes, Petrolisthes rufescens, Dardanus guttatus, Matuta victor, Charybdis annulata, Portunus
pelagicus, Lophozozymus pictor, Trapezia septata, Chasmagnathus subquadratus, and Cryptochirus
coralliodytes, corresponding to isolated, usually old references, never again cited in recent revisions;
Enoplometopus holthuisi and Neopetrolisthes maculatus, cited only in non-taxonomic books; Neoliomera
pubescens, Actaea calculosa, Forestia depressa, Forestia scabra, and Actumnus globulus, for which it is
clearly indicated, in systematic studies, that the revision of this material is necessary; and Trapezia formosa,
re-examined in Paris and perhaps belonging to a new species (P. CASTRO, personal communication).
Furthermore, about 30 additional species, recorded in ecological works, with sometimes only provisional
determinations, could be added to the above mentioned species (cf. for example, Calcinus minutus,
Galathea aff. amamiensis, Liomera laperousei, Paramedaeus simplex, Actaea aff. glandifera, Zozymodes
xanthoides...).
The status of 15 species is doubtful and must be revised. They are: Coenobita carnescens and C.
olivieri, that could respectively be synonyms of C. perlatus and C. spinosus; Galathea latirostris and
Cryptodromia coronata, two species whose exact identity remains to be defined; Thalamita minuscula,
Parapleurophrycoides roseus, and Platyozius perpusillus, described from very small specimens which
could only be the juveniles of more common species; Xanthias tetraodon, possibly a synonym of X.
nitidulus; Etisus punctatus, described only after drawings, the corresponding material being lost; and
Porcellana monolifera, Actaeomorpha alvae, Portunus alexandri, Grapsus depressus, Ptychognathus
crassimanus, and Sesarma jacquinoti, 6 species described from French Polynesia a long time ago, and
never recorded since.
78
BIOGEOGRAPHY
For the Brachyura, FOREST & GUINOT (1962) have already established that the French Polynesian fauna is
a part of the Indo-West Pacific fauna. Located at the eastern limit of this area, French Polynesia is
characterised by a lower diversity than in the Indo-Malaysian area, considered as the origin from where the
Indo-West Pacific fauna has extended. This assumption can be verified here for other groups. For the
Astacidea and Palinuridea, HOLTHUIS (1991) records 27 western-pacific species (zone 71 = Malaysia,
Indonesia, Philippines, New Guinea), collected within the first 100m, against only 14 in French Polynesia.
For the genus Clibanarius, RAHAYU & FOREST (1992) report 20 Indonesian species, against only 7 in this
work. For other diogenids the comparaison with the Indonesian fauna (in RAHAYU, 1992) reveal the
following discrepancies: Aniculus 4 vs 2 species, Calcinus, 23 vs 10 species, Dardanus 10 vs 8 species, and
Diogenes 18 vs 1 species.
In our list, 21 species are known only from French Polynesia. For most of them, it is doubtful that
they are real endemic forms: 3 have been described from very small specimens which could be juveniles of
other species (Parapleurophrycoides roseus, Platozius perpusillus, Thalamita minuscula); 2 are sublittoral
to deep species recently described from material collected with difficulty by the use of deep traps
(Alainodaeus rimatara, Medaeus grandis); and 8 are some of the doubtful species already mentioned
(Coenobita carnescens, Etisus punctatus, Porcellana monilifera, Actaeomorpha alvae, Portunus alexandri,
Grapsus depressus, Ptychognathus crassimanus, and Sesarma jacquinoti). Concerning this last group, let us
recall that Ruppellia granulosa, never recorded since its description from the Marquesas, is here proposed
as a junior synonym of the Lydia annulipes, widely distributed in the Indo-West Pacific. The 8 remaining
species, which could be true endemic forms, are the following: Parribacus holthuisi, Micropagurus
polynesiensis, Nucia rosea, Nursia mimetica, Lissocarcinus elegans, Acanthophrys cristimanus, Ozius
tricarinatus, and Macrophthalmus consobrinus. However, it is likely that some of them are distributed at
least as far as the Western Polynesia, and that they will be recorded there when more collections are
available. As an example, Calcinus nitidus, formerly considered as endemic from Tahiti, has been recently
reported in the Samoa (POUPIN, 1994a).
The French Polynesian fauna could be related to the fauna of the Hawaiian islands, which are of
similar origin and geomorphology. A comparison between the two areas remains difficult because no
detailed list of the Hawaiian fauna is yet available. We however notice that 4 species, Albunea speciosa,
Charybdis hawaiensis, Panopeus pacificus and Sesarma angustifrons, are still known only from these two
areas. ELDREDGE & MILLER (1995) have recently published the number of Hawaiian species, by Infra-
Order. The same calculation, made after our work, is compared with the data of these authors in table 2. The
most obvious result is that the French Polynesian fauna is almost twice as rich as the Hawaiian fauna.
Nevertheless, the fauna of the Hawaiian islands has been well studied, with some important works, like
RATHBUN (1906) or EDMONSON (1959, 1962). This discrepancy could come from the as exhaustive as
possible approach that we have adopted in our compilation. In particular we have included: about 30 species
recorded only in ecological works, with sometimes incomplete or only preliminary determinations; 45
doubtful species (uncertainty about the taxonomic status or the effective presence in French Polynesia); 92
species belonging to the deep fauna, which has been intensively studied and collected recently. Yet, if these
three groups are eliminated from the calculation, the result remains still clearly higher in French Polynesia
(326 species vs 246). Thus, this observation would reveal a real difference between the two areas, the
number of species being greater in French Polynesia. A similar result has been observed for the barnacles
by NEWMAN (1986). This author explains the relatively low diversity of the Hawaiian islands by their great
isolation, in particular if the low islands are excluded from the chart (see opt. cit., fig. 2), and a settlement
from the southern hemisphere, in part from French Polynesia.
79
Table 1 - Number of French Polynesian species, by Infra-Order and Family. The littoral and sublittoral
species come from the above compilation. The deep species have been published in a previous work,
updated here in Appendices 1 (The 12 sublittoral to deep species, listed in both works, are counted with the
littoral and sublittoral species).
INFRA-ORDER FAMILY Littoral and Deep species TOTAL
sublittoral (>100m)
ASTACIDEA Enoplometopidae 1 1 2
& PALINURIDEA Palinuridae 6 3 9
Synaxidae 2 2
Scyllaridae 5 1 6
subtotal 14 3) 19
ANOMURA Coenobitidae 8 8
Diogenidae 31 6 3)
Paguridae 6 1 7
Parapaguridae 10 10
Chirostylidae 2 Dy
Galatheidae 8 19 Di
Porcellanidae 17 17
Albuneidae 1 1
Hippidae 3 3
Lithodidae 1 1
subtotal 74 39 113
BRACHYURA Dromiidae 4 1 5
Homolidae 8 8
Latreillidae 1 1
Dynomenidae 3 1 4
Raninidae 2, 2 4
Poupinidae 1 1
Calappidae 5 2 7
Leucosiidae 4 3 7
Majidae 13 3 16
Parthenopidae 5 2 7
Eumedonidae 1 1
Cancridae 1 1
Geryonidae 2 2
Goneplacidae 3 3
Portunidae 54 1 55
Xanthidae 123 14 137
Trapeziidae 20 20
Pilumnidae 9 9
Carpiliidae 2 2
Menippidae 13 13
Gecarcinidae 5) 5
Grapsidae 35 35
Pinnotheridae 1 1
Ocypodidae 8 8
Cryptochiridae 2 2
Hymenosomatidae 1 1
Incertae Sedis 3 3 6
subtotal SUIS 48 361
TOTAL 401 92 493
80
Table 2 - Total number of species in French Polynesia (this
work, deep species included) and Hawaii (data of
ELDREDGE & MILLER, 1995: 7).
French Polynesia Hawaii
Palinuridae & Astacidae 19 14
Anomura 113 43
Brachyura 361 189
Total 493 246
Few regional distinctions are observed within the French Polynesian Islands. The differences
presented in table 3, where the number of species is calculated by archipelago, mainly indicate differences
in the number of explorations, and amount of collections.
Table 3 - Number of species by archipelago. The number in
parenthesis indicates the species known only from the
corresponding islands.
ARCHIPELAGO TOTAL
Austral 28 ~=s (7)
Gambier 79 (10)
Marquesas 82 (25)
Society 282 (103)
Tuamotu 226 (49)
The Society Islands, with Tahiti, almost inevitable during a stay in French Polynesia, is of course the
best studied place. Nearly as many species are known from the Tuamotu Islands, which illustrates the
importance of the collections made during the voyages of either the US Exploring Expedition (Ahe, Manihi,
Rangiroa, Reao...) or the Albatross (Fakarava, Rangiroa...), and those made by SEURAT (Hao, Marutea
South...), RANSON (Hikueru), or MORRISSON (Raroia). The atolls of this archipelago, without rivers, are
of course not colonized by fresh or brackish water species such as: the Hippidae of the genus Hippa, the
Grapsidae Varuninae (Ptychognathus, Varuna), the Grapsidae Sesarminae (Labuanium, Sesarma), and
some Ocypodidae such as Uca chlorophthalmus (cf. POUPIN, 1994a: 71), Macrophthalmus convexus and
M. consobrinus (POUPIN, in study).
At the southeastern part of Polynesia, the small archipelago of the Gambier Islands, despite its
isolation, has been relatively well sampled, thanks to the collections made by SEURAT during his several
years stay at Mangareva. In comparison, the Marquesas, far more extented, in the vicinity of the equator,
remain poorly known. Affected by particular hydrological conditions, at the origin of the remarkable
absence of a reef barrier, they, nonetheless, do not have any regional characteristics. The 25 species that,
within French Polynesia, are still known only from these islands, are almost always very common in the
Indo-West Pacific. The only exception seems to be the absence of the coconut crab (Birgus latro) whose
presence was never verified during our frequent visits and inquiries.
With only 28 species, the Austral islands have been clearly less sampled than the others. The
northern islands (Maria, Rimatara, Rurutu, Tubuai, and Raevavae) have a fauna similar to the rest of
Polynesia, and a lot of common species, although not yet recorded here, have been observed during our
="
81
stays: Panulirus penicillatus, Coenobita perlatus, Calcinus laevimanus, Pachygrapsus plicatus, Cardisoma
carnifex... and even, in the mouths of the rivers of Raevavae, the big portunid Scylla serrata, common in the
high Society Islands. Far more south, at the southern limit of the tropical area, the island of Rapa, and the
islets of Marotiri, are affected by the particular climatic conditions prevailing in that place. The perceptible
decrease of the water temperature allows only a feeble growth of coral, without a barrier reef. Ashore, the
vegetation is affected by a milder climate, the coconut tree being almost absent. This particular situation has
an influence on the decapod fauna. Some species, very common elsewhere, have never been found after
several visits and inquiries: the coenobite Coenobita perlatus, the coconut crab Birgus latro and the land
crab Cardisoma carnifex. In contrast, at least one species, the lobster Panulirus pascuensis, has settled in
these islands, whereas it is absent in the northern Polynesia.
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nn se eee
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— 1973. — A new species of Decapoda Hippidae: Albunea mariellae nov. sp. from the Banda sea.
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— 1975. — Note additionnelle sur les espéces indo-pacifiques de Quadrella Dana, 1851 (Crustacea,
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94
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95
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ACKNOWLEDGEMENT
The authorities of the Service Mixte de Surveillance Radiologique et Biologique, G. MARTIN and C.
PAYEN, have permitted this research through the collaboration between their Institution and the
Laboratoire de Zoologie des Arthropodes (Muséum national d'Histoire naturelle, Paris), where most of the
literature was found. In the Laboratoire de Zoologie des Arthropodes, A. CROSNIER, J. FOREST, and D.
GUINOT, have always assisted us in our research, and have contributed to improve this work by their
corrections and advice. A.J. BRUCE has helped us for the english translation. Finally, the recent collections
made in French Polynesia have been greatly facilitated by the whole crew of the Marara, and her last two
commanding officers, R. AUDIGIER and M. BENARD. To all of them we wish to express our thanks.
APPENDICES
1 - DEEP SPECIES
(from 100m, and deeper)
With the exception of the shrimps (Dendrobranchiata and Caridea), this list resume the work
presented at the International Senckenberg Symposium, Crustacea Decapoda, Frankfurt, October, 1993
(POUPIN, 1996). Full references on the origin of the material, location and depth of the collections are
available in that work. Some species, collected or described since the first compilation are added (in bold).
For the new species the reader will find more information in the following works: CROSNIER (1995) for
Pleurocolpus boileaui gen. & sp. nov., DAVIE (1995) for Nanocassiope oblonga nov., FOREST (1995) for
the new genera Ciliopagurus and Strigopagurus, GUINOT & RICHER DE FORGES (1995) for the revision
of the homolids and the new genus Yaldwynopsis, HOLTHUIS (1993) for Scyllarus rapanus nov.,
LEMAITRE (1994) for the genus Sympagurus, MANNING (1993) for Chaceon australis nov., POUPIN
(1994b, 1995) for the genera Justitia and Naxioides, POUPIN & MCLAUGHLIN (1996) for Solitariopagurus
sp. nov., and SAINT LAURENT, de & POUPIN (1996) for Eumunida treguieri nov.
Twelve sublittoral to deep species, followed by a "* " are common with the previous list (cf.
Conventions).
INFRA-ORDER ASTACIDEA
FAMILY ENOPLOMETOPIDAE
Hoplometopus gracilipes de Saint Laurent, 1988
96
INFRA-ORDER PALINURIDEA
FAMILY PALINURIDAE
Justitia longimanus (H. Milne Edwards, 1837) *
Justitia vericeli Poupin, 1994
Palinustus unicornutus Berry, 1979 (Coll. 1995, Fangataufa, 250m, det. POUPIN & CHAN).
Puerulus angulatus (Bate, 1888)
FAMILY SYNAXIDAE
Palibythus magnificus Davie, 1990 *
FAMILY SCYLLARIDAE
Scyllarus aurora Holthuis, 1981 *
Scyllarus rapanus Holthuis, 1993
INFRA-ORDER ANOMURA
FAMILY DIOGENIDAE
Bathynarius albicinctus (Alcock, 1905)
Bathynarius pacificus Forest, 1993
Ciliopagurus major Forest, 1995
Ciliopagurus pacificus Forest, 1995
Ciliopagurus plessisi Forest, 1995
Dardanus australis Forest & Morgan, 1991 * (Coll. 1995, Rapa 70-115m, det. FOREST).
Dardanus brachyops Forest, 1962 «
Strigopagurus poupini Forest, 1995
FAMILY PAGURIDAE
Solitariopagurus sp. nov. Poupin & McLaughlin, 1996
FAMILY PARAPAGURIDAE
Strobopagurus cf. gracilipes (A. Milne Edwards, 1891) = S. cf. sibogae in POUPIN 1996 fide
LEMAITRE (1994: 378)
Sympagurus affinis (Henderson, 1888)
Sympagurus boletifer (de Saint Laurent, 1972)
Sympagurus bougainvillei Lemaitre, 1994
Sympagurus dofleini (Balss, 1912)
Sympagurus planimanus (de Saint Laurent, 1972)
Sympagurus poupini Lemaitre, 1994
Sympagurus trispinosus (Balss, 1911)
Sympagurus tuamotu Lemaitre, 1994
Sympagurus wallisi Lemaitre, 1994
FAMILY GALATHEIDAE
Leiogalathea laevirostris (Balss, 1913)
Munida amathea Macpherson & de Saint Laurent, 1991
Munida ducoussoi Macpherson & de Saint Laurent, 1991
Munida evarne Macpherson & de Saint Laurent, 1991
Munida hystrix Macpherson & de Saint Laurent, 1991
Munida lenticularis Macpherson & de Saint Laurent, 1991
Munida longicheles Macpherson & de Saint Laurent, 1991
Munida normani Henderson, 1885
Munida ocellata Macpherson & de Saint Laurent, 1991
of
Munida pasithea Macpherson & de Saint Laurent, 1991
Munida plexaura Macpherson & de Saint Laurent, 1991
Munida polynoe Macpherson & de Saint Laurent, 1991
Munida profunda Macpherson & de Saint Laurent, 1991
Munida pulchra Macpherson & de Saint Laurent, 1991
Munida rubella Macpherson & de Saint Laurent, 1991
Munida rubrovata Macpherson & de Saint Laurent, 1991
Munida sp. cf. pilosimanus Baba, 1969
Munida sp. cf. solae Baba, 1986
Sadayoshia aff. edwardsii Miers, 1884
FAMILY CHIROSTYLIDAE
Eumunida keijii de Saint Laurent & Macpherson, 1990
Eumunida treguieri de Saint Laurent & Poupin, 1996
FAMILY LITHODIDAE
Lithodes megacantha Macpherson, 1991
INFRA-ORDER BRACHYURA
FAMILY DROMIIDAE
Dromia wilsoni (Fulton & Grant, 1902) *
Sphaerodromia ducoussoi McLay, 1991
FAMILY DYNOMENIDAE
Dynomene tanensis Yokoya, 1933 (Coll. 1995, Fangataufa, 310m, det. MCLAY).
FAMILY HOMOLIDAE
Homola ikedai Sakai, 1979 1
Homola orientalis s.1. Henderson, 1888
Homologenus broussei Guinot & Richer de Forges, 1981
Hypsophrys inflata Guinot & Richer de Forges, 1981
Hypsophrys aff. murotoensis Sakai, 1979
Hypsophrys personata Guinot & Richer de Forges, 1981
Moloha aff. majora Kubo, 1936
Yaldwynopsis aff. spinimanus Griffin, 1965
FAMILY LATREILLIIDAE
Latreillia metanesa Williams, 1982.
FAMILY RANINIDAE
Notopoides latus Henderson, 1888
Notosceles chimmonis Bourne, 1922 «
Notosceles viaderi Ward, 1942 rl
FAMILY POUPINIIDAE
Poupinia hirsuta Guinot, 1991
FAMILY CALAPPIDAE
Calappa aff. hepatica (Linné, 1758) = Calappa sp. nov. (GALIL com. pers.)
Mursia hawaiensis Rathbun, 1893
FAMILY LEUCOSIIDAE
Oreotlos encymus Tan & Ng, 1993
98
Oreotlos potanus Tan & Ng, 1993
Randallia serenei Richer de Forges, 1983
FAMILY MAJIDAE
Cyrtomaia ihlei Guinot & Richer de Forges, 1982
Naxioides teatui Poupin, 1995
Naxioides vaitahu Poupin, 1995
FAMILY PARTHENOPIDAE
Parthenope (Platylambrus) poupini Garth, 1993
Parthenope (Platylambrus) stellata Rathbun, 1906
FAMILY CANCRIDAE
Platepistoma balssii (Zarenkov, 1990)
FAMILY GERYONIDAE
Chaceon australis Manning, 1993
Chaceon poupini Manning, 1992
FAMILY GONEPLACIDAE
Carcinoplax aff. cooki Rathbun, 1906
Carcinoplax aff. crosnieri Guinot & Richer de Forges, 1981
Carcinoplax aff. verdensis Rathbun, 1914
FAMILY PORTUNIDAE
Charybdis paucidentata A. Milne Edwards, 1861 *
Parathranites hexagonum Rathbun, 1906 (Coll. Eiao, Vanavana; 155-240m, det. Moosa)
Portunus nipponensis Sakai, 1938 *
Thalamita macrospinifera Rathbun, 1911 *
FAMILY XANTHIDAE
Alainodaeus akiaki Davie, 1993
Alainodaeus rimatara Davie, 1993 *
Banareia fatuhiva Davie, 1993
Demania garthi Guinot & Richer de Forges, 1981
Demania mortenseni (Odhner, 1925)
Epistocavea mururoa Davie, 1993
Euryozius danielae Davie, 1993
Hypocolpus mararae Crosnier, 1991
Lophozozymus bertonciniae Guinot & Richer de Forges, 1981
Medaeus grandis Davie, 1993 *
Meractaea tafai Davie, 1993
Meriola rufomaculata Davie, 1993
Nanocassiope oblongaDavie, 1995
Paraxanthodes polynesiensis Davie, 1993
Rata tuamotense Davie, 1993
Pleurocolpus boileaui Crosnier, 1995
XANTHOIDEA JNCERTAE SEDIS
Beuroisia manquenei Guinot & Richer de Forges, 1981
Mathildella maxima Guinot & Richer de Forges, 1981
Progeryon mararae Guinot & Richer de Forges, 1981
99
2 - PARTIAL IDENTIFICATIONS
These incomplete references were generally found in ecological works. To avoid partial
identifications in the main list, they are presented separately hereafter, by alphabetical order. Four genera
are cited for the first time in the area: Glabropilumnus, Heteropanope, Libinia, and Tylodiplax.
Actaea sp.
Actumnus sp.
Dromia sp.
Euxanthus sp.
Glabropilumnus sp. :
Heteropanope sp.
Libinia sp.
Lybia sp.
Neoliomera sp.
Pachygrapsus sp.
Paramedaeus sp.
Parthenope sp.
Petrolishtes spp.
Phymodius sp.
Pilodius sp.
Pilumnus spp.
Tylodiplax sp.
Xanthias sp.
MONTEFORTE, 1984: 170, annex 1, tab. a (Moorea, Tahiti); 1987: 8 (Moorea).
MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea). — GUINOT, 1985: 452 (List).
ODINETZ, 1983: 208, with a ? (Tahiti). — GUINOT, 1985: 448 (List).
PEYROT-CLAUSADE, 1977a, annex of the species: 26 (Moorea), juvenile; 1977b:
212 (Moorea). — GUINOT, 1985: 450 (List).
MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva). — DELESALLE, 1985: 289
(Mataiva). — GUINOT, 1985: 452 (List).
PEYROT-CLAUSADE, 1989: 113 (Tikehau).
SENDLER, 1923: 40 (Tahiti).
PEYROT-CLAUSADE, 1977a, annex of the species: 27, juvenile (Moorea).
PEYROT-CLAUSADE, 1977a, annex of the species: 27, juvenile (Moorea).
HOLTHUIS, 1953: 32 (Raroia).
PEYROT-CLAUSADE, 1989: 113 (Tikehau).
MONTEFORTE, 1984: 174, annex 1, tab. a (Takapoto). — SALVAT & RICHARD,
1985: 350 (Takapoto).
MONTEFORTE, 1984: 173. — PEYROT-CLAUSADE: 1977: 25.
PEYROT-CLAUSADE, 1977a, annex of the species: 27, juvenile (Moorea).
PEYROT-CLAUSADE, 1977a, annex of the species: 27, juvenile (Moorea).
MONTEFORTE, 1984: 171, annex 1, tab. a, photo p. 131a (Takapoto); 1987: 9
(Moorea). — SALVAT & RICHARD, 1985: 350 (Takapoto).
THOMASSIN et al., 1982: 393 (Moorea). — GUINOT, 1985: 453 (List).
PEYROT-CLAUSADE, 1989: 112, 115 (Moorea, Tikehau).
100
INDEX
A Gffine: PENCHONG. cosiaso cored reece cece eee 71
GODT eVIGIUMP Cr CnOn ew neater ease 71 affinis
abbreviatus, ACANIROPUS ...se.sssesssessssssessessssseesseeeses 71 DCLG sian ctssesnssteaiaasetsinvsnagecnes sussesas oosenanserueenerne 55
aberrans, Garthiella, Pilodius ....c...cc+c.+cssc+sssses00se000- 54 TNE LTT ION CA tes pee PAP eee 5)3)
Acanthophrys Cristim@nu ...cc.cccscsssssssesssesssseeeees 26; 78 Galathea wanes. dais ayant athitd ae 19
Acanthopus PON CHOM ooo ee che 71
GQDDIEVIGIUS ares oo sce canee eee eases see eee 71 SYMPAQUIUS -.eossseerressnesecssaseeeernsnseceesnnseceennnsesetins 96
DIGTUSSIMUSTs FOO RA os soca tact chee emenctee coteck 71 AKiaKt, AlAiNOdAeUS ........-ssseeeresseenevneeessneecesieecnsneseenns 98
LORUUTONS sion vsesaastocticacocnessndtituns wusatiinia, antares 71 Alainodaeus
Achelous, Neptunus, POrtunus ....cc...cc0.-sscs00ss0e000sse-- 31 RIOR, che Meccssnscsnnancnatons cctansn chee cee on inaten cone eae 98
Actaea VUNG Gone cere ee eee sass stacdnns exes Soees 7; 41; 78; 98
CUT LIBLS eee oes sh a ee ae ee ge 55 albicinctus, Bathynaris ........ss.0errsveerseeresseeerenveeesien 96
COICUIOSG see mic tisseet coe ete ee ee re 42;77 albolineatus, GrAapSUs......ccrieecreeerieer viernes ceenneeeens 67
EGV IPOS EO RUSS, Tete ee eee can 44 AlbUrned SPECIOSA .......eeseresvneseeernnveeercnnveeeeenns 22; 76; 78
GOMSODI INR cece co er eee ee ROO 42; 45 AIbUS, PSCUdOQIAPSUS .....2-.+s.csseeroseeevneeesaseesasesenneeeons 69
Ganges eo 42 alcocki
EDI ESSENSE aie RO ee 43 LiOXAntHOdES ......seeeeerervineseeerrrenveseeecnnnneeseeseennneee 49
GQ GTTOUE a8 acon sti sce cs cactcsci cece ke RR 43 TRALAMit ....seseesseessseesssesceeeseeseseesueeseseneesansennes 36
SEITE CEPT ATO ble PESO teria Bis Shit 42;77 alexandri
HUSUUSSING een er ee 42 Caine ctes wooseeeeeeesesnseseeeessnnneeceennnnneecccennnassee 31
Renee te 2 al 45 er eee ee eee, 31; 77; 78
DGRV Uae ere Ftc reer ee ne 38 Algae, GAlathed «.....eeresevsvvveeeeeeeeseensnnniisteeeeccennenninss 20
POLWACONINGR een en one e 42 alvae, ACLACOMOTPHA ......r.e1vseseeeeeeneesninns 5; 28; 77; 78
FUfOPUClAIGs WAREMAN 3s chen lig SP ena 43; 44 Amamiensis, GalAHed ..........sseerreeerreessnreerrsveevsees L977
PILDPeLIOI eS wh erat weeks Rested. SEE: 42; 45 mathe, MUnida see sietr fev scesensusenasinsosaseossontacentoeete 96
SCD Gi eer eM te recrets Mece cha sects ceete atcha 43 Amphiucd, UCO viesssssesseeseeeeeeeseececcrecerreseerennnneiesieeie 74
SPicee OO Te 99 Gnaglypia PlatypOdi@, .oerecnxcc-cnecn-necnnecvennenensoreaeeeeer 46
SUPER CUUGT Ss seat cin tee ai oes ease ence 43 Anaglyptus, ELiSUS .....000.ereeereecreerrneereerneeees 6; 52; 53
LOMENIOSG cre oer Me eee res 43 anceps, Eupagurus, PAgurixus ......--crseerrrreeenneseensees 19
tum lOSat eT AEE BONIS ho oes 44 ANA COSSY1, CYMO ....seesseesvescsesseeeseeeseesereneesnceeeenesnness 38
Actaeodes ANQUIAUST PUCTULUS?. Jc foteceess- +20 sssccatente tooeeseseeteeetere 96
OPTI S oan ee wae ee Se Ne re 55 angustifrons
GREOIEES oo ate tse ee ce 42 MicipPOidess ..........-.-ssveseerssseeseesoseenesnneeseennnsceenen 27
CONSODI INS Pics Semen. ee i, em ee 42 SCSAPMA ooessservssesvesvecrseecnssecsneccnesennesnseeenees 71; 78
Hi SULSSUMUS Tht Oe ek ee 42 Aniculus
FICHIERS a ein ae ce ee anne: 4] CAIAIGLULLGS YRS, Bo Reavis cc. csi socscusessanssaetcereees 13
FONIETLOSTIS cr eT ee et 43 TAAXIMUS Bascave coksouns ons oaatisot ince swan eSeeaeee Re ee 14
Actaeomorpha SPs osvdeahateeeteaeess Petese ee once ce nsoanacs neat eee 14
AVaelteeti on ees in werent enn 5; 28: 77; 78 EY PIOUS ss ic ee cue aiwccoussncxsseateuas atone inane iateettarereceeeee 13
COS octdor erates etree Nene Sa ae 28 aniculus, ANicUlUs, PAQUrUS ..........0rs.erreeceveerrseesenees 13
DUNICIAL Doe cxsan cae coe RAG 28 annulata
Actumnus GRAN Y DAISK. 85 BS Sieh cake Biss ccssncasnce sbossseonsseleee ee 30; 77
SPORE ee PR A CNS Ia a ce 61 GOMLOSOMA........0eoseeevveeesnrenseesnnessneenseennnceniennneennes 30
DONUT access coe tecseon niet tests oes OTs 61 annulipes
Aigitalis. cccsscvcicosesssss: A ae 61 Lid... essrsessseecoseeeshecesseseneesneeesnesnseesnnenseeennes 64; 78
GIODUIUS 3. eee ee ee 61:77 Ridp ellie xs. cccchecccct cote cecs co teectea cee tevomnene nents 64
INLEGITIINUS ecard toon ee 54 antarcticus
ODOSUS Riss, Geetha ee TR ha Foster 61 PAPTiDACUS «...000sessseessseeessvensserseeessnerssseesssees LO; HT
SCLC sce snsins disse De ee 61 SCYLATUS ...sesccssseeessssssseseneecnsneensnnesssnnennsneesanees 10
Sis snssanvonatiodsveasess al atsaee Meee TR CEN RR 99 Antipodar um, Ar Clides ..........s.-crsveerrseeerneernneeennesennnee 10
fOMUNIOSUS Ah ps See E PR A RO ace scisis 61 Arctides
aculeata: GalaMed i ciccsnncen ie eee 19 ANU POAT UM ose esesssesecessseecsvivessnnneesnneeccnnneeenneeen 10
Qdactyla; HIDDG wnas. te. acnvteaea en eaetie nore 23 TOQAIIS .oesssssseersseseecnssneeenssnsscssnecesnnaseeennes 10; 76; 77
admete, Thdlamita (.it NALA NRW 33 GCUALUS, XANENO ..ererserresvvvrvvrrernnseseeeenseceeeeeeeneeeeecen 48
Geneus, ZOTMUS ec. ake Le 47 areolata
Gequabilis, CLDANALUS ....sccsesssescsessseessessessssesseeesees 16 CHOP OG OSES cos. ci veicintiesnonscuskeconassvacosencoueceraneeeets 5S
A Cth SCTUPOSG «o cscsea-tts-nireanctasetnn eee 28; 77 TrQPeZiG sssveesseeeeeveeveveeseersssnsnssnnnnnnnencscees 3; 58; 60
areolatus
A CLACOdES eee Net th cscseccstzniensvies RR NS 42
ILO Shed Sots osscareoceuccsisntie nee OR 55
GVTNU CH NN OCA PUOES | <-ccsc0zc-csocsscn-vsesisteseencorsetesee 54
Ashtoret
PV ATUIOS Cres meticectvaevsersevesceves scives mesesttens ee 25; 76
LUN GI IS be F ces ccke tevb hetviseesisvessesass std et ee ee 25
DICE fed Se vedateh deh varias teceasee sussa recceninn dd Peeters tees 25
GASP EN HACIUTIANRUS | occ ccccs su sesus cco cavesesussseoraceen teense ees 61
ASPET. SCHIZOPNUGYS orcs sensed acoceassscoae-Jesesaeeseecoees 27; 77
Atergatis
CLE GQ GNS cssarmscsce Scien CR ee ee 46
i LONI AUS psscucoecesieeevaucvnveeeesesteis SO ee 45
LEN DGLUS x tesescscchas cssedessessstens EMO tes ase 47
OGY OC fae icv sis satan taausanteccorssnent sxsunciesbeneerostere 45
Atergatopsis
PONG Eases rscatbstracatvsersrectscssessssesia See ote 45
SEDNIGLUS yecteessssis dca ssesiwsscuctcnsesstivectaseseesez ete 45
Aulacolambrus hoplonotus ......10.ccssseseveeseseseretenees 29
GUT OG YSGY UGiUS <.:ccach-css02csscseesesenssiscucesttoneees 7; 11; 96
australis
(CRACE OT os eI es 95; 98
DYES sp oec see aca ee eRe 17; 76; 96
B
BGISSTUSIPL AICP ISL OMe as tetera secss tenses ase tce eee 98
Banareia
GUI os sho soos ssi sc sedenateaduse she nuscdis saute ee 98
DGIVILIG* Ceiein oct eee wa ssecocciemen tania 38
DQNSKESM QU ava aanene cote ee Tees 25
barbata, Chlorodiella .........ceccsceccceseescessenseteeteeteeeens 53
BON fi} CONE CF os crac stecsencsscscestsuse estas Wuseessesso 11
Bathynarius
CGIDICINCLUS nextevcccevtussxassiesiseits 96
DACIPICUS here ncesis SIERO 96
bella
TAO Giascehoscsccetststcsauetsesatssats Be oes 38
DY AD OD zest ewistvasvicsesvateertsxs teh eo eee 3; 58; 60
bellus, Actaeodes, Carpilodes ..........:cccsccsesseeneeeeee 38
bertonciniae, LOPhOZOZYMUS ........seeceesesesesereeeeneees 98
BeUrOiSid MANQUENEL .0.......seeeseseecesesceseneneteceenenceaeeens 98
Didentatus, LAChNOPOMUS .0.........cseseceeserseseseneceenetenees 48
DifFOntalisHELiSUS cee S152
SETI STOTT 0) Nee OSE ORCC COLOR CORO EG 11; 80; 81
DISPINOSUSN PEI Olisthes iver eres eee eee: 21
Doileaui, PIEUrOCOIPUS ......cccsccecessesseesecseeseesceseeeee 95; 98
Doletifer,, SY MPGQUIUS «0x2 scsscseesssicesentteceoeesanteestonne ete 96
OMNI er yA CLUIMNUS csvescisesteusesstesdiersivescecives Seer 61
borradailei
TI YASt NUS sss sessccscsssessvzesasresseva Treen 27
BelrOlisthes xccocsctrsrtiesiti scan Sees essences QUEVAZ
bose; etrolisthes: 2 ks ees 21
bougainvillei, Sy MPAQurrus .........eccscecereseseeresenseeneee 96
BOUV IEF URGIGIULG ereecsessssesencossceseceentssussssrsesth ree 33
brachyops, Dardanus .0.......cccccescesevesseseseneesene 7; 17; 96
brevimanns;| GOEnODIUG vcccccc.coocecscx tote nto eessteesiees 12
DIOUSSCI TLOMOLOBZENUS nsec. -cvecessa-oerentesderaccesnceteoetes= 97
101
C
CACSTILEr ONLY DIG icsacccntascsccssscsescestscs detect ne Seo 37
Calappa
COILED P GN scersescsverwssedeesvewestbeussatbehe bees teettbees 6; 25
EDUCA ie sae v asst sare cinarsscesavecesosscenscacesncevece 25; 97
SP ANOV Sacsesussaievavchswadesteceletvasades caste dssaessStieee reas 97
LUDENCUIGD sede se ese se eee Ds)
Calcinus
CLE GIS hese dt oso ee sevens dccsostisseitees OR Ne 14
SD GUNG. A iets weve sc srscndesel sates ssesseeseoet nase ee teete oes 14
ID MCMCNISI Se ee clos oc tec xiesress sestesssestesocesnosttes 14; 76
ER DSLR Roeder ae 14; 15
UN DONIGIS iscszstvecs sesesteeccasscsseseee casa ceeciv ROE 14; 76
IGEVIMGTIYUS od ee eee AS 14; 81
IGLCnS css cece eek IIS)
TRINULUS eer s, Ee De 6; 15; 77
ALERT TTS cpooz asot oe REEL EEOC EOE eee 4; 15; 78
SOUL ate neo k Mosca aes kateatstscaseeenunee ee 4,15
SDIGQUUS 25 cvs cscwescbescicucasassceviswestiepesesuiien Meteor 4; 15
LCV AE TO DINAL foi. ccstsesosconsusestzsh dodveetsneetescse srsbeee IES)
LIDICEN crore sok oot an Soci eeete osm See co 14
COIGUIOSQNACIGCG ree hh ee 42;77
G@allinectes|alexand rips rescsccsccccsasnsstoveonctescesteeesen se Sill
CGMPOSCI GI CLUSA aire eieccsretteescsesoievess nes oreo eet 26
canaliculata, CryptodrOmi@.............1c1ccssersesseeseeseees 45}
Canaliculatus, XANtKhiAS ......0.ccccccccescececesceeesseccesseeeeses 50
Cancer
GIFT oe eesctas i sees cecsrei statis Sa 11
TARA DRS OE CEES EEC DEE EOS Ee econ octoaos 10
Caphyra
ROMP ONS Sed vast teats svastsaseccveteestesesseenttees 29
UL UOIS Risse ech aie Ae See 30
Carcinoplax
COOK IB rece store aa savectta ed eoad tate east eases Woaewe ne 98
CH OST ON Uta eerare eae sc cieacdsvsveuetsosec niece ss Oreo 98
WEN ACTS TS yo tere ea osc aco aewsucaasbsaedoct Sees ROMO 98
Cardisoma
COMPUT Os seins bessescks tani tas tee veases tees 65; 81
TELL POS eave. vescsdet covaceesc cov dates stsatazecesets eee eset 65
OD OSU a as cides oasis eed fess ck ores Siem es eo 65
VOLUN UINE reotenaes se eo pan nace 66
CArinipes, ZOZYMOAES ...........csesecseseescsssseessesensetseees 47
CAFNESCENS, COCNODIA ........cccceseccscceescsseeeeaes MDE TTS Ths
CArNIfEX, CAP AISOMGA ........esecseserseveesesceseeeecsesenees 65; 81
Carpilius
COTIVOXUS iF ess ech oS SR 7; 62
TILAGUL GUUS aeons ain aoc a Sea Season 62
Carpilodes
DELL SW ees eactate Ga neces ee ein es 38
CUIGLIN ANUS oS ee hs is as 3 55a 39
PT QIU ALUS ecscecrccie wateenctscessesdvwv tk cccsscbeoueteuerotieettes 40
TROMMICUIOSUS Soe occcccccece sas cassn neces des eee 39; 40
QIU AUS os tecressuvcec cass cis suscs savesscssivss sevesessssasseve stort 39
TU OCLUS Ho rxcnccse cos sacs cinctets csstntsstn.dvaxesioe: Meee MOOR 4O,
LIUSLES Meericee ste aces saa aac ane Soe RUIER ate sensu ee eee 40
VEAL PILI CAILULS scat cose one seoeaestvas Soest bene ceeshessanso te es 38
VEN OS US Pesce swakcsswabt evcavesttnccivvseussesbosee ever neato ye 40
GCarpiloxanthUs TUgipes sovissscssssscisesecsecissevseeseccsonenrses, IO
102
Carupa
VGOVIUSC UNG coz ascucsstssceseeveesssxetuanscistexssusnisnieosties 29
LEVIUN POS) coor iae ccteacz seostensrccotsset ee entaceh ore care 29
Catapaguroides
PRO QUS 5 csc verse ssivsusvascsnstasscsstcnttnsetesstenesveseostvess 5; 18
UFICHOPHENAUNUS ees teeter eee 19
CQLOPIFUs NUIDUS: nc cssencsncceseseeissnncasinrsottheue LN? 29
cavatus, Cycloxanthops, NeoxanthopS .........:.1.01000 50
COVIMMANG, DEM GUA. oe rnecsaccatscsnstentes ctnccns- ee rat 37,
cavipes
A CLOL ron vaccas echlcrigatcancsasecteonees eonsocnascaneas te 44
GOCNOBUG cei cscsrase ee lo 77.
TSQUINUUS iii sscsccecteceutevsseatsete cgentssseintestes Sees RE 44
GAYSUUS PSCUAOZIUS ccrtescccscsanescccnascessceaness eee 65
CELLOS PSQUITIS Orne eet ge 44
Cenobita
GOI TRESCONS .o vanysas gos acs ers acstetissicsien essence ee 12
GLY CAG h.nss.n.enscosssocevarsssescecatencteysen Mey ayie 12
OUI CTU oe covaccascqeas sie cuesestgsanezarsocesees CATERED 13
TUQOSG orn rissa. sesstvessusasscesecaesessacnese seein SEES 13
cerathophthalma, OCYyPOde ........c.ccssecesersseseerevensenees 73
Chaceon
CLUSU AUIS rine ere er 95, 98
OUD UL, cis sasocrncseunponsesusste changers sas test ee 98
Chaptalt, TRGIGMIUG’ cio. cccsccsccsccsscocses entre 33
Charybdis
CETL LG oor cw ncrcsecscteskens etesce nce ee 30; 77
CF YUY OAACEYIG « ......ccsocaocaseo--rsrsstaaneseteeccospestactine 30
RGWICVISIS’ . 5.5 o.csccss-steccct ies oeensnee eR 30; 78
ONVENEALIS 002 Vi acecinns auteur a Ret estes 31
POUCIGENIGIA.. .. avcsectsconcnae em Cota 7; 31; 98
Chasmagnathus SUBQUGArAtUs .........ccceeceeeesenseees 69; 77
Chimmonis, NOtosceles ........00...ccss0cccceserscecesessceees 24; 97
Chlorodiella
DOF DOUG e.lcbec cans teste rte asset TR 53
CYR CRED sero nevis cscivtensoseeenss sos: csceucss MORSE EOE 33)
TQGVISSHIIIG .ccted Uvccaccatters seciueies ascot enet oa ERTS 53)
LQEVISSIINUS.. cssssue cencsseasortccnsses hu tosee+0es SSUES 54
TGC oon scecctuactanccnien ersten sac teeehe uae eee 54
Chlorodius
CYTE OG wossvecssssssoesessnsiaecssansvesspscssscesseS SUReE RNR 216)
CONGAAW vcccssececsu cess Seneetxene to ceeeet tac cnie sen cc ME 55
LGC VISSIINUS). ».cscessszscssessspsadsvacesceseconssnes sate tere as 53
INONUUCUIOSUS) ic ccssesscectesssseseriaa vac cess oR S13)
VU QED. cvcsscseeas caxsssises scessciansvsacsoereninsesdesces SOROOINGS 54
SGMPUINCUS) ccryspceconssasseareteenusssenssonsns MASE es 49
UP QULALUS). vc ocecccsescxeansessaryrac Meee eee eee 55
Chlorodopsis
I COLUG sis scsccssnssssiconsecesnssrsis siete tent ee entaaes 55
QT ANUIGLUS oo scistesccssocceessvetore ceonsssvestaevec eos Gee 56
PUUL; 10s scossivasess sivseepsaccansauaesssecensveczssscee eg MNteyRNi 56
SCODIICUIG ccc ccxcdarxsacaanasnstanaisarerieseniis SO OES 5)
SDUUDES.. cssccessacvacvactrssstscascvaccssveccecsatvccsts BRYN’ 56
VENUS senpisescisainrscestnpieabsbancncscssertoesescste tee 57
chlorophthalmus
AINDHLUCG socereseersossnvssussovenaseaeevess «1040 ORO LEEERO ASE « 74
GETGSIAAUS Sooo sn, <ccccucs cag scnnntenssesecets Ae 74
0 677 epee RR ie i RE aire Ta 74; 80
Ciliopagurus
KU GIND IL on ces teorettsscesessstusseg cargo tt eG 16
TIRGJOM, svronereoccsvoesores sopanesssapeeiasrxarsr Sieh Ne RRR 96
DOCIICHS crs cssesossescssarce-cistecessecrsisvasesss 96
PULOSSISE,secsocossncossoegyses sors: soes 2PNGQUEGs GAARA 96
SIP IDOLUS iv csoavevixseaseucaus saves conceccest eect eedscee Wace eeenee 16
CINCLUNANG: LAOME OS coe ee 39
CINCLIMGANUS, Car PIOUS oo scs-2n-cosavsexieos acts sosese tes 39
Cifactipes; Vein apias. enctncesseczsrtaccsusssaveneesactestet eee af
Clibanarius
CE QUGDIIS, .csnnscniecsstsennestesaettcrncesosarosSRteieeT ei 16
GOT GLIUAUS <ccnes canta saan ioakanstcaante sc saan ass 16
CUTY SLOT TUS, |. .5-.02-<nconesnansnnnen-teiee i eetpeees Ne 16
RUMILIS cose oe ee 16
TAPS ONL ca nsssopsiicadsscoosdbospoosencssncevedsevan St R ERE 5; 16
PRADA OAGCIYLUS ssassnsnssosvasesnncosessasetseosacess De 16
SCTE I ALUS acts cans nsec cms dacksaasnosaskiunceeee ss 16
SIV LOLQUUS .cdoatsacadaasinaacenansdanssaonaananass ote tote aaa 17
ZEDI.G wc sadenApadavnndinkispasnasscousaaoa. ee 16; 17
CLYDEGIG: | C CNODUG . ...2...0s0500s000ss0ven0s00sen0saeeeu-0+0 ee 12
clypeatus, Cenobita, COCNODILG........ccceceeceesecsecetseee 12
COCCINEG, POL CCN ANG vosasccersresacsoancpaassnonacos eae Pal
GOCCIREUS: PCI OLSTRES <..cccs0ceseszeunssess<¢ds-s Se 21
Coenobita
BrEVUNGIUS csscsecessnsosesso0ssssoussesovusessesses date te ae 12
(AHA ROT) epee ee 12; 77; 78
GOV IOS a sack secs S ile cat cnaetn nsesiaiuseeee eee Sel 2077,
OLY PCCLUS 1 cnonenrnennanconaneneGhtheaeaeas Sis aaa 12
BELZENRGON SE .vssessscsiseresconoscosonsyorenes baeeeip aekeoesa ae 12
OUD CN Bacay ts ace toewdaee ee 213 dg,
POETIC. cesssusasseesasensstsseseaceosneexsseasasennse eee 13
PCI LQUUS sce cnsisaaas nica c 2a anl np pnahoe ata L2 24392; SI
FULD OSUS -. ncuncocncsncsuoissaseasenacosnevacec-seeses-t aoe 13
TUSOSUS VF. CTANULATUS).....-<-n0-.020--n00-scccscacewsseats 13
SDILOSUS is ics ssnnsssiucctidenacselacestauace Cae 122 13477:
GOCKULI POSS TP HGIQMUUG . 2cccc.cnens-desastenesncsntessn-2s 7 eee 33
consobrina
UNCUT A) Nepean e 42:45
AGCIUCOD OS® O88 Ses ceeacen sak secctestaeace ee 42
consobrinus
A CLACOD ES) ike Be cocnece eigen an one'coe svsnesesceiss see ee 42
IMGCrODEELGIMIUS . ot ccccc0ccccecs <cs-0senssc ota 72; 78; 80
Contrarius, Parthenope ..............<bsveswsresstpac te 28
convexus
GT ee 7; 62
MGCrOphiha linus. ...r...i0..s+sn-.n00+es0000-4:venseetoe 73; 80
convictor, Eumedon, Eumedonus ............01.ss000-+000000+ 29
COOK IC GV GINOPIAK iorvn ccs secs osacsncescenssneacccueneeeene
COOPER 1, TRGLAMNG ....<:.01s:seseeeseecnscse0cesss ERAS 33
COFGBIRUS: ClIDANGTIUS 5. occcc 0c cvececsxckccsouen sce 16
coralliodytes, CryptOChir us .......cc.ccscseeeseesereeess 74;77
Coralliogalathed Rumilis ......cccccccccccseeecesenseesecseeseees 19
COT AUMGNG, OCY POE. ..5.-..5550:0sn<-- gaetanatagtint- nga 73
coronata,Cryptodromid.......<cstgserehet ee 233 77
COFTUBALG, TRGIAMIULA..........:0.00.0.000.00 arneSechs-e pees J; 34
crassimanus
FrePlOdtus soos sssssvessicasn seve nee gbtctbromamt eaters 49
MaGCromedGeus ...<..0..0:000-00: SH 49
Pty GCRORNALRUS. ...o..c.cssconss0000- kus 69; 77; 78
DC EE Cs ee 49
CHASSIDOS). CQ os scvessesenevosesenas scseneee suse nisi CNG 74
CRASSUIN, SGT «15.000 50sass0usscaesso sence seaaaeeetangeenee 70
CEO. TRA AIG soc cocasduvccays tess coxseesas eRe 34
crenulatus, Hemigrapsus, Heterograpsus ......0.00.0.0 69
Ghinipes, GEOLLADSUS. ..........aaavicpyscgseyprns- HEM 66
cristatus
eC PTOULUS oo. s0ysycsaissenqesesieasess Ree eames 48
LOphozO2 yu ...vsosso0ssss0. aortas) Re 46
RGHINOGIUS «cccicvscccsssnsnces irene ign Gee 47
cristimanus
A COMINOPIIY S. csccse ac seccvsivscexssus chess sscss Been 26; 78
AY ASICNUS 2 costestcocreteeses mame 26
Grosilerv,|\CarGinOplaninvccscc eee eee 98
Cryptochirus Coralliodytes .......0cccccceeesecesserees 74;77
Cryptodromia
CANA CULAR RIS ss avsccariveioss ist toevcssviarunsateer tees 23
COT OMG GEREN INNS oa is ssa Saves case ene eeme aac 23;77
i fLLLIZE Got SR: atop ee ou eee 23
CryptodromiopSts tridens .......cccceccecersecsvsensssosseseeeees 24
CUDULIFEHMPOLVACCLUS cet icce crete eee eee nea 37
CY ANCUS PIGNES A csesteteccetsecsesce cette cccceae cece 68; 76
Gyelasisuborbicular sie seccccecee cee eee 26
(Gy ClOGChelOuUsy POMUUUS oreo enna he: eee 31
Cyclodius
IDV AGI US yevk des shoe toed eae vevaae devo edatededcaaeeeca see oe S15)
ONNALUS a Os, save asvabs suis cataraees seme ett wok 55
Cyclograpsus
FLED Cre NS TRE Dc orouasecce otawougeuaom aaa eae: 70
LONG IDES ccc aee vice civ oechex sit osecctes soy scraease ace aa dsc 70
DOT VULLUS He oss cc sat cacteeatd oa cazswiavadacecc tin eieeetatabees 70
Cymo
GMANCOSSY Ui covered osece sev iret noberrueereistecoe ro tata 38
EPL ANGLUS reise risk a aa secs eae tees agtete dace ee seta 38
MMELANOGACIYIG .cecescnsencsuesngusest saan naraten sep Eesti 38
MelanOdachy lus iecucy renee Behera ere oe ale 38
QUGCRIIODGLUS scission ee 38
GYMOAOCE) TT ADEZIG ere cersccesceneeneecesenees 58, 59; 60; 61
GY RENAL) OUGANEL Ge. be vcs cecess cts Seve setei cee eisas ee ass S7/
Gy lOmal Gillet esses cerctdesecs vote tees ceess eee EEE 98
cytherea, Chlorodiella, ChIOrodius .........0ccccccceseees 53
D
Dacryopilumnus Cremitd ........ccceccccseeeesetetceeeneneceeees 63
IDR RA REL eeepc neers e oe Rae ree ea Tee 75
AGKINDMUNGIGMILG mre ees 34
DAA Or fiatnorrid deen enn rere te 28
danae
ACLALG eastern er Rte nr eee 42
TLL GEO RD ITI TALE 3470)
GANIEIAEWE MIN OZLUS mnt tne re ee een 98
Dardanus
GUST. CLS HE erties Sneee erie an bennett: 17; 76; 96
DRAG T iy Op seeriter naar ks cess tenner settthe Mee ies 7; 17; 96
defOrmiSen ene aes ater Er eee 17
QETIINGL US ist MsntnnesEseneersrhPeSNeTTE RIT eset certaeaererts 17
ULL GLUS ts NG NEE ONE EERE ness 17; 77
AGT ae eats aera TP ey ner eee ee Uae 18
LAQOPOdES Re Nes NE Ren Ee 17
TILED ISLOS reteset tances rent ene Ee re oeee tts treece 18
DEAUNCLIALUS RIOR Me er tee teeettert teres teeters 18
CHCA TO Direccrcterse rane cracerenstater tou cae eae 18
SCULCLICLILS Here seieiitenties staan ronmental Ve tee 18
davaoensis
JEG ONT TS tccoreecpetcence Hot rete eee RCTS ID 48
ERED CZI Ge nee HOA NE etc: 60
GECACanihiswPetrolisthesimncs cee 21
deformis, Dardanus, Pagurus .......cccccccecccssesseseeseees NY
Achaaninehlorodiuswermneee ne 5)
DCC CO Le 74
103
demani
ESLESUS eee sees cece ccecutscs sus stcsecuesasecuvagevecessoess- eee 52
INCOM OME Gee ceditseteieciset ecchh oa ote 40; 41
LAE ANTI eee PORTED ROR ETER ERECTA DR ees 34
Demania
OR esate ns ccasin sce ates cani Siesssxapcttataeiscs BAER: 98
TOT LENS CNM i wien eacee tic tu ca ae seis Seea cba astasie Re 98
CHIGIA SI ADEZIG ane eee een 58
DeNIGtUS EUSUS) viccsscccccesssacsacesscunsencescee tt 52
Geplanalisy Gymorecsene seers eae 38
depressa
TA CLC ore cei sccundaceacknsiav ces ecu sicsnteasncteh sieve MOMS: 43
OG ESL Gi ereancse eyes sae ciax ctocedendosuecasesssecssia OM 43;77
PlAQUSIG IEE aka eek REO AID 72
epressus, GraPSUS .....sccesececcesceesssecsesesseeses 67; 77; 78
AUFOMMISHPGQUTUS Ko iies ck. cce1.- toe etree oes ete 17
digitalis
VA CLUITATIUS oot eeu was eenaa ane gndse sesame suszanss SoC oes 61
IRIGY POdIG ca eestisct ees BE 61
ROD OZN A ivvssxceccesvscecesieessivteesesttenassieest ee RAI: 59
GIGitalisMl i Qpeziabereee eee ene 58; 60
Diogenes Qairdineri ..........csccccceccsecscercesceseeseesecneeseess 18
Discoplax long ipesy ci vcevcsvsresecxcoutesetucusvs wee tees: 66
distinguendus
IMGGhOMmedGeus Ee 49:77
MAINO ve ccditeee eis He NEN NEN VEINS 49
dodone, LOPhOZOZYMUS ........ccecceeseseetecseeneeseeseeseetenens 46
dofleini, SYMPAQurus ......cccceccsecscsceescesessenseeteceeeenees 96
Domecia
LGD ee reel a EO 63
IS Pid eee irs ae eater ere ete 63
Dotlla fenestrae acct. 76
Dromia
ISP si stn eia ss oun vcben Lisareedneveaueestocsdsecsecvessses A, 99
WEIS ONDE Sh Atle se ince cece 7; 24; 97
dubtusW#Portunus s4h ieee chi Be. 3]
ducoussoi
Mind diesen china 96
SPhaerOdrOmiaenncenci sie 97
dumerrilit, TylOCArCinus ........ccccceseeseeseeseeeeesceeeeeceseeees 28
duperreyt, GELASIMUS ........cccceccceccecceceseesseneenseeeeeee 3; 74
dussumieri, Gelasimus, UCQ ..........ccccccseeeseeseeeeeee 74; 76
Dynomene
hispid Gsihicniadeccsaiiste ee. OR 24
DV GEU Al Officaseestccrcssrosescactentetseeen eee 24
SELES) as ter teen renee ren ee occasions ssansaeuse cic 24; 76
SP SPER swececniseesesoassetaceanativsvaveacriacacee MEER 24
SDINOSQIATE sccrsied secissavivasicosissvcerasazes RR 24
MOMENSUS srescds, Nasexscccheasscssesiahasasesed sasdaa cons Oooo 97
E
easteranus, PtyChognathus .......ccccccecreeerecees 6; 69
EDGLIG’CrOSG = eee ene ee 26
ESDALIOPSISICFOSA een teens 26
EChinO€Cus PentAQONUsS .........scccecceesecscesesscesersenseeseeee 29
edwardsi
EL OPROZOZY MUS vars beececceeteee acter 46
SGAGY OSRIG ss scescs ssvssssticcsesscesteassisestscisste 97
TRGIGHUO’ A esc cccmeweideanmsi I. 34
104
Eldredge, PetrolistheS .........:cccsccscseerceccesersercenseenees 21
electra
EMSOES ieccverehic ein ee OS. 52
IE LUISUS ERA sic ciicensiocstteecitvaaccsscee 52
elegans
AL ER BAUS HE sv cdivtiuesdcasivstecansvieeeneeee ee 46
CGIGIRUS AR cacecccsiseeteccviaeswereves deers SORE 14
LiSSOCQLCINUS ........0s.cceccseesecescesectsevteazenten 5; 30; 78
POMOLISINGS i. cccsscscesecsasecsssest.nesccss tts ee 21
ENCYMUS, OLOMOS .....ceeseccesessesseesessesecsscesecesscseceseens 97
Enoplometopus
ROUPUUS ES sevussvectreccoacavsssasiveccecisessventteen ae 8; 77
SP ANOVA sssasuseousscseessvevcocssevascnctavceessexetie ee eee 8
EpigrGpsus Pouitus ........cccsccssccesscssssesseseseseees 4; 66; 70
EPiStOCAVEA MUSUTOG 0.......ssessecsesevserseessnsesecneseesensons 98
eremita, DacryopilUmnu .......s.ccsecsseceressevenseseeseeees 63
Eriphia
LGEVIMONG esse ssscceer essed isieiesssse ORE 63
SCADVICUIG sxticseisiessistestsxace seins ROR 63
SODONG sc vavccedensed borestaces saseisoavaeeiveed tarts SO 63
erosa
ACLACOMOPPH| .....eeeecescesssseensesstsneenscesenseeseenesenens 28
EDQUQUE:. ccseccssescivascaecseccxansnrtses BAR Be 26
ELD QUIOPSIS esicesescorseececseverssoicccovensssevsdsstee POE 26
erythrodactyla, CHArybdis ...........:.:cssecseerserceseseceeeees 30
erythrodactylum, GONIOSOMG ..............sssseecsesensesees 30
Etisodes
CCC GER ivvccs eect tie ROE 52
PLONIGUSE. Bic cwedscsestivesi insist eeien ata ee 52
SPLCNIAUS seuccsscossadvciesuclivessianaccctetiarieccte see. 53
Etisus
GnaglyPlus: 28 IER ecctshadesccee 6; 52; 53
DIUfFONIGS BEA i oscecscitiscasts Beccustot eae 51; 52
DOTA Rica ssn 8 cosa tescteassans canes saeascvuasaannddenaessOiees 52
CEN GEUS is cvevzessaseucecatuaeagevashiciosteats eeatste TR 52
CECI sis swavencascgestieaxnateedndeeax cae SORE 52
JROMUUISU RE cos at cseccccsec czcstn inten dsoeves eeeveioce MERE 52
LGOVIMNGNUS orc vavsascssinascseeveveescesdee secnestsaeetenes 52; 53
MACK OD GCLY US cass coane ac us snacuceee seas 52, 53
DUTICIGLUS | cssacesseciietecsssesece. note eters: 53; 77; 78
FUG OSUS )e vsceiccusseed suvans i van scexmeteeananera nee wees 52
SPICNAIGUS .scsccccsevuscoucouymntsesshsseteeaeer a apie teaRe 53
EWCH GLC RR SAA vicccsa cds veszavsaee sabia tice nase ERR 75
Eudora tetr QOd on... ...cecscscccsssssssessersesensesesseecnenscnsesenes Sl
EumedOn CONVICHON u.....secsesesseseesccencenceenscsecnsenseaecnsens 29
Eumedonus
CONVICIOS casscccicaceecetsnnsssivansans tastes Bete A 29
POPUL GG ONUS isos susesnsscncsvievacedeacenssssicesces BRON» 29
Eumunida
RED RAE swstaneinuier auntie Renee 97
WO BUL OR: ccsvaceucsasuee veveseeduei den saeveddaccevethecoteeanete 95; 97
CHOPSIS) PAQUIUS i .sccnsscaynsencecoescesesedscivossssessuss (ossesesesn 18
Eupagurus
ONCEDS 8. vcvssswssssocwsivannseteovoaentaboveseuceseneeteese dit averees 19
NGEVINANUSYS,. csssssescsonicstucuteen eines nearness 19
VAQOTUSRAisceosiecnsinsevseseswnesinctuivus Reet RAEN 19
EUr YOTIUS AAMNEIGE. cvccssssvsvavsvessnvavasesssoieundacousecseepnsness 98
CUryStErNus, CLIDANALIUS .....cccsesceeerererreeeeseeerseeeneeeees 16
Euxanthus
OXSCUIDIUS sxosssannnscasessansssssndeaiasaas taba eaten eRPEBSS 41
FUQOSUS coicsissesestecsacvsceies corageeqeenaisnce ROMO 41
SCUIDLILIS. cs scsivesscsseinscedrcrrccnchanensce eee es 4]
SPAS ones ova sonassnsvansoxascanvesades acts tv giguoA teenth Calvan 99
CVAINE \MURIA .ivecsuccsaseviassicescovsrtevsrvaivateonsietenesecs tenes 96
exaratus
TLC PIOUS esc. hactsei ved sascatiadsodsssivss Me Oe 48
MANOVA AINE 49
excelsus, PETiQrAPSUS .........cscscccecessecsseeeceenseeteeeecesees 65
OXCENITICA, PAFACIACA........ccsesceeevecccececccccececeeceeseseeces 43
exsculptus, EUXANthuS ...........cscccccssseseescessessesctsessens 41
F
fakaravensis, PAChYQrapsus ...........1..scsceseesseseee 4; 68
fallax, CryptOdrOmid .......cccccccescseeseeecseeseeeceeseeecneene 23
fasciatus, PANUTITUS 00.0... cceceececceeceseeseeseceesecnseeeeaes 10
fatuhiva, Banari 00... cesses cette eeeteeseeseeseeeseese 98
femoristriga, Panulirus, SEn@X .......1.ccsccssesceecseeserscsees 9
PONESUG) DOUG onc cosccccescsecchecesuessenevacassocesecacseesenette 76
ferrugined, Trapezia .......s.scccsssesessesees 58; 59; 60; 61
flavopunctata, TrAPeZia .........scccsecsseserecssesenscneeseneees 59
PLQVUS SPUOGIUS fo o- cn c.cenccverectoocenesoxencsusccersacesce ee meee 56
PLOTIAUSTATET SALIS (iio... 0-c cane coscscceeosceccensroossesesortenscnens 45
Forestia
DEPT OSSG ceva scssiscsvavesesousedess savattceriescvetseeoctem 43°F7,
SCOOT rence et EET 43,77
SOTMOSA, TVAPCZIA ........eecseeecseceeseneecesecseceveecseeeees SI 7L
fragilis, CatapagurOides ...........ccceeceeeecescesceeees 5; 18
frontalis
FLUUSOD OS veo o ieee clseaeccenne vas secon ctacesteessnazkacsseecerncenee 52
PEVISUS) fos ctesicsa.zevotcsorersaesentvetensceseuneevteeses eee 52
PUSCOMT TAD OTIA Gertie stucttntottceonceat ncaa ee ae 59
G
Gaillardiellus
TUCP POND pi AE ccs poss ses cas wsecasavsseceseaattscsoctwane 43;77
SU PON GULIGTIS PINEP wsec soto nttercscducenasnscucemes seen 43
gaimardi
CGICINUS .. acid csscesesysoctess signs cenyerenegeene ee 14
GEL ASIINUS 25 Gis oins suis oni cescescnsenenh-h ccane gee 74
TCG. EePB EA sede HERI acess sneentennne pee ee 74
QAirdineri, DiOQeneS .......c.scseseeseceessesersessesecnseseecseees 18
Galathea
CCUICAL BASS Rene onncnen cin casevatntesnducOee tania 19
OF AIS issesbok Hak ti tease dete aaven eeneen dere cee 19
ee ee 20
AIVAMNIENS IS) oo cco sc oneve can weven usec eaeneeceny 19;77
LAUT OSENISE. RRR oven doo vseravenedostouese urns 20; 77
MEQGIOCHIN AE Sioa cen wcevonsnstons sonsenssver neuen 19
SEDI OSUPIS vs svseasseesn vonesececseendsvevensvsssecvestuaueeneeenes 20
Gardineri, TRALAMItA..........ccecceccesseseesecseceeeseeseneeearers 36
IPA EUL A CIACDIOR Sos c0cniscacenssceppicviento eee ier 43
FETAL II OVA TTC Ron Sep Ree eee eee cao ee a 98
Gr thiella, QDELT ANS, ..cc.c.cascossesovsoseverasarascesycoanetenanse 54
gatavakensis, Thalamita .......ccscccsccsesccseeseeeeneeeees 4; 34
Gelasimus
CHIDLODRIRGUAUS .occcosiessnnowcxsosvensnecetee PETRIE 74
GUDELT EYE CER B hs a Raat as 3; 74
AUSSUTETE RR Sc wsnsceascvaoncenattinenieeneyreicaamaeseete 74
PALMAR AU oxincecvivacescusyasinsconen shasta eee Ses ReneS 74
LAINE AWE sersicnvscsssonryusr enna oun ennrnt eh eantean 74
PulohellUsy misc vsrcossercrsosses cones teatyereteaean tenn 74
LEI AQ ONION wissesseeiecnsoveneunecsensasecern enlaces CARELURERCEENE 74
REMAASNUCIA sisccovtscasncieoeswesercateetee hatemeeene eee 26
i
gemmatus, Dardanus, Pagurus .......ccseeeserereieies 17
Geograpsus
GIADA DES ccoaeceesececpeocecsoncbocenoduenrosacoaposoccocodtooCBeuCICCaS 66
SEPTATE coca asceostnececnacak cachissstaascedso eso ceoaadecccboosesdiBacb0c8 66
LGV ACL TS ierreictecnoct tenes concsadsastemersrereverser soe 66
SLOTIN IN ar een al stakeasesersvonesmbaaeat tect oresevarens 66
BEFMAINI, ALCL QALOPSIS ........ecerercseeseseesnseseserecsererenes 45
glaber, LOPROZOZYMUS ........c.cecesesesseseeenes cess enecenees 46
Glaberrima, Tetraid .....ccccccccceecsssecsessessesensenesenees 57
Blabrda, DOMECIA ..........ccsceescesetees ene cteeseescseesesenenecnees 63
Glabropilumnu Sp. ...ccccceseccesesccsvscsetssersescetecsesesens 99
Blandiferd, ACLACA ..........cccccrseeseeseses ence esereteneenes 42;77
Globopilumnus QlODOSUS .......ccceeseeeesseeetes esses 3; 64
globosus
GIObDOpIUMNUS 0.00... sceccecesseses cre esetete tenses eneceeees 3; 64
PULA ES ee Ue ere teens 56; 64
Globulus, ACtUMNUS ......cccccseecsee esse cee esetetenseeenees 61;77
IONICHSISWN GIGI G)sstens certs enestan-acereosncccageteetoteees: 34
Gonioinfradens, ChArybdis .........10cccssccesseeeeereees 31
Goniosoma
CAIN ALUN eee ener re nescence ene neeteetineceess 30
CV YthrOMACtYIUM........sceeseceecessescesensesensesesscaeescneees 30
LENICAL TIT RIS con cer oceee roc tecevetceeve cactiaesniat sores 36
Goniosupradens, Charybdis .......0.0cscsceeeresererees 30
gracilipes
GAPS USE NI NRE a eer eaeues aes Ntatere 67
EIA aR Re RINE aE Me Coeteesseeteoeeterece 35
gracilipes, StrODOPAQUrUs ........cccsvccesecsccrecessererersesens 96
gracilis
Cy CLOGS REAR NN Te sn teeaaee coer 55
Ee PEOCIUS ENE resto cescs sess ete teat sceetestresssseestets 49
WV IOCANCITUS Te een c Serres ts secce ess eouecnrcagescoaeeete 28
ATT OREN I nae Ne loeb cersevnersonagsecsestontsatiess 49
BrAcillimus, GrAPSUS .......csccvecceesssessesesssssesnseeeesenees 67
gracillipes, HOplOMetOPuUs...........ccccescerssecscrssereeserens 95
BVANAIS, MEdACUS ..........csceecsecsescnsssesseneeeenes 41; 78; 98
granosimana, Liomera, Pseudoliomera................0. 44
QIANOSO-MANUS, XANLNOES .........0..cececsssscrssssssseeees 50
granulariS, MEtaS@SArMG .............eccesscsescscsseesssssscneees 70
granulatus
Gr PilOdes ier ye eee cecnececedssntetecsostossescnsescteats 40
EhIOROdOPSIS Boccia hee N ee ectotaps tetcdehessceseoocbeoseens 56
(COLON Sere iRe oI CEO DDE OE OBE 13
INGDUUNIUS rea II ola teadBecestentaseancsstessetvess 31
BU MOd Is MR ie sccctvescsctstecsdmeectetgts «eeemseo rs: 55
PORE UTLU Sartre eet tote ne se ren sacsccussetarscetessecr 31
granulosa
IAS TOL EIA ERO eee ieee eae PN aS 25; 76
LOPUGCICON nese tose cteseece te oescuet vtiattaetenes 47
PLA DOdIG Se Se aN nPeneietcieotsese 47
RUD Pelliaire ee secseneseos ssteasetvouesaseticsicewesesss sees 64; 78
BraNulOsus, PHYMOALUS..........0...ssseseereersseeseneessseesees 54
Grapsus
GIDOlINGAtUS AE ON eee toreenederne tote 67
EPH ESSUS HME SOR UNL DOE: 67, 77; 78
SACD ESO RN Man tonne tenstadecousste sietseesesnarsesereestes 67
AGUS elses aba cctasais teakos cies vestteobteetoeveeetert se 67
IT OPSUS Maree. Neencvncceeseoscaretesviswssce sve treasetsae ston evs 67
LOWS IGP SISO IA. eee oateccnteelcieelonseserteonotesteies 67
TRACULALUS ee caeeetsnesetecsssretsrgeneeseresesscstesesstes 67
IGUUSIemneit elo etetentencraceteecenstststeveccecsscctrsuescessear 67
SEI OSUS reer coronas ces Necsus asickeueeatsvhectsecsenseetoaes 67
LMU CHUSTALUS) BU ctccsulsscocsncsreroccecoesassssrepsseorears 67
QIAYE, GEOQTAPSUS <0. c.iiecencenssscocenseesoseneucsoconsatonsesses 66
105
QUAMENSIS, CAICINUS .0.........eccecerseecentecssseecetseeeees 14; 76
guinotae
IPErEnOn eee em ikecs licens LM RM 71; 76
IPORLUNUS PRD Ho eee Ie a Rn eee 4,31
OU GLD MUI AD EZIG eee Reyes sieke secs etc nescocetaas rs 58; 59
guttatus
GV AAS ROR! Pere ed oe aie tsar: th Ye 17; 77
PAB UT US Oe ee NON SU ct aetee stones 17
H
RAAn UD GRA ANUS Mee ins cscesreieetcessseesueoraessonssvexsveetarse 18
Hapalocarcinus Marsuptalis ........00cccccreieresteeiee 74
harmsi, Liocarpilodes, PilOdius .............0c.0ccccceeees 54
harpax, ThalassOgrapsus .........cccccccceccerecreseieess 69; 77
hawaiensis
Chary Dds Perr creer career estecsenet 30; 78
VEL StL ORRIN AEM 1! NOEN OO NL estos leresese dese: 97
OZ iis ee NEONATE EL escoet caeevec secs 64
Hellenus, POrtunus .0......csccescessecsscusesseeseesseesseneeseees 32
Hemigrapsus Crenulatus .........ccccccceceeseceseteee essences 69
THAD, scroceneneenecoccosocne es cnn059 0 s0cccaconnESForccNess65000-009C 28
hepatica, Calappd...........scssesessesersessvssesenseessees 25; 97
1 BUG RTTTS sexe pececoccren pec Hore ae obo HOR SC SAbO ERODE URE ESOC CEE 28
herbstit, CAICINUS .......ccccccccceecseceesenseceecensnsceceeceees 14; 15
heterodactyla, Tetrauid .........ccccccecescsseceeeessesceenees 57
Heterograpsus Crenulatus ........0sccecesevserserensereseees 69
HeterOnucia VENUStG 0.2... ccsceecescessessenscesesenseseeseessenes 26
HeterOpanope SP. ......s.ssvcsssvsecsvseseseseesenecsenscseescnecseaes 99
hexagonum, Parathranites ........c.ccccccerersererieeees 98
hilgendorfi, COENODILA ........cece severities senteteeeeeeeees 12
Hippa
GUILT SUA) corse po) po enrscoceos oe beae a0 3 coe Dok NTI FoIODE ENED 23
OV AU SVS Te iy hed sees Aescuiers A REE Se 3}
[UDTER TED cocericcoceee0c03 92000602202 0480209 x50 TOGISEHEO TOG SBoC003N. 23
SD ee ae Deena eak anton senees cretereroteseees 23
hippocrepicd, PAVActAe .......ccccccessserccssrrenssnrerseseees 43
PUP SULAWEOUD INU lara ncestecetcwonceceesioeccenss<eneereersanneters 97
Wir sutissirna, AClAC.......c.cceceercensessecnscescsesesecesssaeenes 42
hirsutissimus, ACtACOdES .........ceecescvsesscsessesseneeenes 42
hirtipes
Car disOmay reece dsectecscscsvanosn ah enon evteaaes 65
TA CIY GAYA cp rcscecceecnecoo CoO eID ODEO EET HOBIE. 58
hispida
DONEC Gi srestersscevs these steshess eaters 63
Dy NOMene vice Reciecticrcsrrnte eR mts 24
holthuisi
EOP IOMELOD US) see racys as oxy ev -cvavestuveuceretesenceaces 8; 77
IRAP IDACUS ee Staenaeloecewateeasses ustceae oes 5, 11; 78
Moma rus, PANulirs .........c.sceeccsesseessesescesseeeeceeneeeeeess 9
Homola
EKO CUE lec cecbocecchnacsalavetses desks SMBS 97
CTA ALO reece ees tree cBae ESSEC CCP EOEE Reon oe SchtccecbCoA 97
Homologenus br OuUSS€: ........1.ccsccccesssseesenessesenssnennenens 97
Hoplometopus gracilipes ..........ccscccecesecscessresecseneenes 95
hoplonotus, Aulacolambrus, Parthenope ...........00+ 28
horrifia, Daldorfia, Parthenope ..........secsereceiree 28
FAUENIAI PH OLEUSiecoossencac-peeteatrcsmesescosesncece settee eee Pais
humilis
GQUIBGNGIIUS oscccchceosocosecs ee eee te ee es 16
CORGINO BGI Ate dieicivevseancooucrad UN ete eee MO
106
Hyastenus
BOTT GAG OD caves cescissiis ccesee se ences ctse roccsaetiensviayeraes 27
GHISUUTONUS |v cccencecsssessstas sot ovevcevassoxyoucstuapuagreee ess 26
hydrodromus, POLAMON .......ccceccsseesessessesssesesesseeees 77
HyPOCOIPUS MALATE uo..csecsecescessecsssnsceceeseesscescteceseears 98
Hypsophrys
USL QED oi os vcnc sus can seucexse sevass toss sirvesverssterstcuiga noses 97
TNUPOLOCNSIS: .vcccccvccersteceucdessssstusssisectvisesensentene 97
DETSOMQUG cscncecsreus tinct cove soc certe wast eee cower ereeieenesce 97
VASO IK, AIG Goo nass cosenec-tocs ence ee taser cass caecaeess vices tceetaee 96
I
PRICE, (CY TOMNGIG vetncivossusth aude Men vasdt uals coop cals stone 98
PREC AP HOMO L scscaccuscesssnccwssoseceucontusniceced cone iseaets 97
Immaculata, PlAQUSIA .........:scccseecesecssescreeseetenee 72; 76
WNDENMAlis, CGICWIBUS oo evccccescscenaesceseeeeceeseesoeetaheceee 14;76
INCISUS, LOPHOZOZYMUS ..........cescesesneveereeececneenees 46; 76
Hap ELV SOP [ay Simecterceccneceezsestecessoentece eee 97
InSilanis, NCONMOMENG .....2......00:-n+ssenesenstnse tis: uc: 40; 77
Integer, CYCIOQFAPSUS .........sccssencseeseseceeeesetecsenenceeees 70
integerrimus, Actumnus, Liocarpilodes................+-+ 54
Integra, TRALAMUA ........cceecceccsesssenssscessssenseeseaesess Ses
intermedius, PtyChognathus .........0.cc1css-cscecceeerseseetes 69
WV GE) POPLURUS |. 5 .vascc-nsescecsseasesusoneonteepoites et ae 5; 32
J
JACQUINOL 1, SESALNA oi. .sceeeeeeeensenseeneenseteeneees 71; 77; 78
TONESIUS UTIUNQUICUTAIUS .0....0..ceccsesensessesesesenetscneneens S/
Justitia
TON GUM ANUS oi cece ceouisiated svat aiecese een 9; 96
VONICOL Bass cco cease svtvass wlsscessis sects consosedasuas tae Noes kes 96
Tuxtaxanthias tetrdodom .........ccccceseseseeserseseceeceensess 5
K
RED EUIMUN dG issen. occ sccvs vas schaeancsccnnessgeeen teeters 97
Kraussia
WGN GUCSGAS: cavcccbesesasegescectesnsusctuderie si satersevek ee SO 5]
FQSUAD OS eviccssvesvcusetencseseivueskori sus seqeansuas eater Sl
Krempfi,\GuiOpG Gurus. ..cscccecceusscccssssu+1.- ssveneeteneeaies 16
L
Labuanium
FOUN GLUT av cvciecsvesdcuaeaucaiustedendedéaysae ROO 70; 76
INGDOZOIMEUIN vas cecssscnosszconsesstheents RANE 70
Lachnopodus
Didentatus need. LAR, PIR 48
POMADENSIS, ..ss.r0n0.0. HTT ARON TES 48
SUDAGUEUS vss ssussssvesvanenseanssconss aavconsnss RAMEN 48
LABUONSIS .cicshisonssassesseseisecuspagesescece sane ere 48
laevigata, Pinnotherelid .......ccsccsceseseeeceseeeessecseescneeee 72
LAEVIMNANG, EVI PHUG ...ncensiosarnssosncn0 eS ae 63
laevimanus
COICINUS ee 14; 81
EUSUS RR ree nn ree 52; 53
EDOQUIUS ro cccssscrcstceseatss ters tere teres tires restr tetscaeer 19
PQQUIIMUSE ei cosccal cocediesetoretestinel sel ctisiresiceectee tee 19
laevirostris, LetOgalathed .........ccceccecsccsecsscecseceesesseess 96
laevis
PH OMEN, GE cesevoncaveaivel ox cauers vs dl obscene sacar eee 39
IENSSOGONCINU Stevsovecveesdeccees aetna ee ae ea 30
OCS POD Cian cosiccnscet scscnsoucnczernsastudionv soc taetee ee em 73
laevissima, Chlorodielld.....cccccccccssessecescessecescennseseees 53
laevissimus, Chlorodiella, Chlorodius .......2...2.0.00+0++ 53
laeviusculd, CArUpa .......sceccsececenseneeeseessenececeacetseeneess 29
la gopodes DAVAGHUS .,...c.+--sn-sosecvecesnvssnssovsnes-taessents 17
lamarcki
Retr OSI Nees ete SEES soeaes was tsveecweper mes 21
DEUTER, i55 eee ne 50
lamelligera, Parthenope ........::sscseeseeseseesecseseenecseeee 28
laperousel, LIOMETG .........1cceceseesensevseeseeneeeenseeee 39: 7h,
lata
INGO RU ce SE RE EO OO: 45
[ERD 2 ANE RT TR EE 39
PSCUd NOME G sszicecsessscazeieseesseessessarece teen eee 44
LatensMEGICINUS: <écc.sivesccousxieiseeiacss kee see 15
latifrOns) XQQUMOS «ives ccverisinsesseveesexestmereest ree 50
latinostris | Galated ers ie tieststeeeeeeceeee ee 20; 77
latreillei, GElaSIMUS. ........1..c:00eseeseecececccececseeneeceecences 74
EatreliGimelanes a veis sccccaaevecassaceacsssatsueovte-haseetors 97
LAUT CORBI QUS orcs cx metezwes Srimeete cee eee 11; 80; 81
GUUS NOLOPOIES) 2. .c.eevs sc vcscrstecccorsecassecsn danse tou eee 97
laysanitt, TWeedield \... .c....cc:snsssts sscsnactrees oseunticee ea ee 2¥/
Leiogalathed laevir OStris .........c.cccccesceseeseeeecuecesensense’ 96
Lenticular is MUNI AG s.ccccs0sciss<s0s00200050050020000soagenaer ee 96
LeptOChelis, LY DIG wc. (vc. <002+.n0tseescs0sap paprtnnse SNe sane saan 37
Leptodius
GUASSIMIONUS ox vcs san coonnssiausseaske-osCaaascesstaa ck ee 49
CT USUML US occa Sia oan BN waco as eSB oa cena Uae 48
CGY GOENSIS oes sists sccsnscSonapetnse nese cus ee eee eee 48
OCXAU QUUSE, fo. scsaccv'scs concusiccotel bacscasnasac tates See ee 48
QT ACLIS ss oscse sc sz sassoesceveeweesisscestraiesososcsy svsasnse eee 49
LCPOD OM oak ccc wes cnceescsece ste densetsssucennn scp ea ee 48
SAPO MIN CUS yo soscvans coos seontvte coceccvent an ae 49
Leptodor EEPlOdiUsy .cccisccsssc ccs cs coos secessenes so tens aoe 48
LeptOSrGPSUs VANE QALUS ..........-<e0c-osses-0ss-nssnanncssanasee 67
leschenaudti, Thelphusa ...........ccceceeseeeserseeeesenseeseees 77
LEWINSOMNI QUGUIENG .....scc.saccssvsswasssacesesssannasnssqandite 57
ENDIVE SIDR aches ccnssat sac onc cusvcovesoansccvnccetctsesssGepa eae 99
TEED YSTOS|UFTCATLTI ONS 5 .c cece. csatn cnc cceonncnessts een 29
Hem Data LOD ROMICIDDG ...nsc.ox..cxncs--neoncenstenenennsetanets 27
UINDGIUS ALCL QQUS . <...0.0.ncsecacnssnsnceatocteesagttannce eee 47
LE ALIN (GORIOSOMIG.. vas «ccscrvasvsacovesonasusasnanienaceenaneee 36
Liocarpilodes
IND Opa re. Sas casos cosnsscaecnenstcarnivesasteerescsetaenteies 54
TRAD IBS bess eh cocetrekaevsnsssacessnks htousacs ouster 54
PPL CDEP TIES ck rccicenk = cancsues soncunscnscuseeponmesgen ste 54
SPD marin cneoneccnan connec sccceck Conte ee teantss Une Gana 64
Liomera
LL I eee et RE. 38
CURGHINGIOR csc es sccce acon a 39
RU ARO SVEN. sco ccsucecsneinc oeceeicese here aaa 44
LER OS eee Ee A fe EE 39
LOD OL OUNC Rica 5h cs concn sacs sacesy peskeerecen beeen an 39° 7¥,
2) te POI eon i A RRB AER HR LE ESS, os ho 39
THRONE CULOSG << oxerencscccensvenseenvou nodes asa nena ees 39
MU
Ipallidarswaunncs. wince, MUNI RN 39
TECHLEN Site reese cts ind aniuencenes ooh, ETS RN 4]
TLL DAY Seo Soa cs calealc vans casey nacaeesuaevsssaicveneasese teeter 39
TUBAL A eine ie ns teasnen ie Sto adss Reea ees 38; 39
PU DEPE Sian ierscie tiation oinutetiase ites waht tts MIN 38
SCMUGSTANOS Ane ae 40
SPIN PSONP Siew sessans tien sontvaccescess Cea ee 40
ET ESULS) eooeeee reheat at cok aad TEE od 3; 40
VENOS GRR vases teen ee eats hasten Sate cee 40
oxanthodesialcochipe ese 49
Lissocarcinus
IEG GSR RI EEE RUS ehanhh earths En 5; 30; 78
TRUS cece Sasa RS ESSEC CEPR ee OEP STONE 30
ON DI CULGI TS re AON ORNL D EIN Tet eM eter eet 30
EIEROG ESTER ACAMING wee nets ernce etre eee nrn et 97
Up LLCy ALA AVI GIUM CR nooo reine 6; 69
lividus
Galeinusi te RO A eee et. Is)
(GOON APSUS 52 vessseiscssiiees ccsseetosestaee iste 66
1 PEREIRA) Ky cher oRCeEEREH eC ESC SS CURE ER LORE He RO ceca 15
LODALUSNOZIUS eT Re ee, oe 65
longicheleswMilnidare nse) anne mee ote e ts 96
lONGICOWNISHPMUMNUS Ser at ee ee eee 61
LONPIMANUSSIUSULICnnct eee ee eee 9: 96
longipes
Gyclograpsussn ier Seek eet Senet eae 70
DD ISCOP LAN ee ee ect ectncterttcta ecietsehe secrete sce 66
BAG BHAD SUS osc cca rscasnsietedtss enc tieceessneeeecaeeee 68
PACU Sra etree oe elton te eee ae 9
longispinosus, Neptunus, POrtunus .......0..0cc1ccccccereens 32
LONBILGTSISN GL GDSUS Ess cence ett neste tctneee steer 67
EOP IACIC ORD AIULOS tree net tenner mrereer esters 47
COP ROMIGIDDAIIMDAIG acetren ecw eke tenes cee 27
Lophozozymus
DEFLIONCINIAC ee 98
CILISTICTUT RY conmcoo peer cece oce roe aes See RE BOER RENCE ECeaY 46
DUDA? crt PF CaO COIS HOTELES POTOSI BOSD 46
CAWGNd Sheree a FS 46
LAD EF esr terorrcccenccscercetceneee ite ON 46
AICS US settee t ci ches ae scant ce STEED WORRY 8 46; 76
OCLOAENIAUS| A Eee eee 46
IDICLOTEM Xchenns cides vet ncwe nes erie chests MN Re Oe 46;77
ISP eMeeheola nse ares as oge ONIN Moen geehoccttenerene sees 47
ISUPENDUS 2. senesxosisaveaescsnsanessccaei vine cyecaazsscsiaete 5; 46
lunarissAShtloretcocnninwe ee 25
Lupocyclus quinquedentatus .........cccccccccsccssessersecsess 31
Lybia
COPS ERA eerste veccneirnnnnaucce 37
LEDIOCRELIS ev cccscstesesereesoumentavewcdvirees 37,
(PLUM OS Oirrnrercciue cent te 37
SPlditecavstsnsdstsvateecverservaeevindtdvaaadossstratevrsss SOMO 99
LESSEN AA reccrcscccstrisrectrarseiecsvsceciets MANORS OM 37
Bydiaiannulipes iinadtecnctaniie Mee ees 64; 78
M
IACTOCEN GN OGNDOGEN ir it ere mce ATE anne te 76
TIACRODAGIYIUSHEDISUS Harte ttts eterna eee BydO ss}
Macromedaeus
CV GSSUTIGIULS Drm area Tames WERn eee, 49
CISHINSUENAT Ss Renna creek eres 49; 77
107
EUCLID OS stv diadrencter cies ta rae DEMURE OO ON TS Sy, 50
Macrophthalmus
GONSODLINUS eee ee 72; 78; 80
CONV OXU Sie ste iasessens aeeciomsedS/ieec Dee a 73, 80
DQIVIMOANUS etrccrtastte tinisese catetotiecs ee ek 72
SCENE isweee eaten iumstaiatthsibatnavie cers oases eee 73
macrophthalmus, POrtunus ..........1cceccecceereeeeeeee 32; 76
MACFOPUS, TRALAMIULA .........creceeecceeceecesenteneteeesees 35; 76
macrospiniferd, Thalamita ............cscccseseereseeee 35; 98
MACUIALA MEN GD CZ disc eciecterecesiee tt eter eee 61
maculatus
Garpiliisscmsiscessetnccsie se: EE RE ae 62
(Grapsusimopretin iscsi nraniaiiinsen AS ea 67
INCOpetnOlisthese ee eee ee hence 20; 77
MAGCHIOSGOUGCTEll aw er ee eee 57; 76
magnificus, PAlibythus..........cccccccceeeseceeneees 7; 10; 96
MAjOPNGIOPASUGUS\ en srseecet sree cec eee ee 96
majonayMoloharecw sec a 97
MANGUENEL I BEULOISIGlaxesrescctseres sss gsceennet eee eee 98
maorus, Eupagurus, PAQurixus ......1...cccccceeeseereeee 19
mararae
Ly POCOIPUS oxcichee sa esedestviececuecnctessuescssoseasSeteesetterees 98
PHOGEMY ON ssseeaticmaces cizitiesstceeesesusiesces Roh 98
margaritatus, PIUMNUS ..........1..cescereeceseeeeere tees 54; 64
Mangan tiferanMIGipparenwnew nnn eet eee ee 27
marquesas
IK GUS SU ca een seh shite cers indies ends Meee ol
IRalapediamiciitesin uit nis acer Le bo Sl
marsuptalis, HapQlocarcinus ......1....cscceccceseeseeeeenees 74
MAU GeiETQMmen ary eewn xs meuln ss: per ssenes ALON le Ts)
Meathildellannaxi mare en ase se see 98
Matuta
DTS KS iiseenstart nepal ents oak Ota tadnnn elo 2) BY 73)
DICT aie eises steep eateries uss pynmandndnsocintst srs Na Eee 25
VICI reenter eta, ME BSS UY.
maxima, Mathildella ...........cccsccscsssesecececcccsececcseseneeaes 98
IMOKIMUSVANICULUS sesesseetc tazeseasicetcesstassessssisetneecetees 14
Medaeus
DTN IS eavenaemte sa nerassseticol mo Wisststee esr) 41; 78; 98
NOCIENSIS ORC ncn! ACE EN Seer 42,49
MME LACANUNGAMUROA CSN eteres cc sict econ contseeesseercaete see 97
NCR AIOGR IGN GAARA reteesattocetee teense ete cnret 19
NEBESLOS DD) NA GNUS cecere st ecceetnc ess cceacesecetovecstneteeteeices 18
MELANOdACHYIANC VIMO a. cecanclocsesc ce csccnscersostsis can sneesoee 38
melanodactylus;\ Cy MO weccveississcesnssoicesse sass sites ossnssees ne 38
IMCL Gite SSCLAL A) Sais ceca ccdinvasenvens Svcscedecsnteseebescatesd 3,
Menaethius
TONOCEN OS assoc Bietebessvevtesecestee te ea oot Ro 27,
TUDOR GUIGUUS ee Servis sess ccass stvcsscs te Sesiscaxscess MOE 27
NG ARATOTT RON LA 71 Terence Pe eee Ree EO ES ES 98
IME RIOL A ULOMAGUIAL As scenece coceewcccrece-neee tee 98
METOACNIALUS) PAIUIMMNUS) wiscesssss-ssd ioe neem ene 61
MESSON,, /MCLOPORT. GDSUS)..u2c+.c2n0.->00.00ce-csecnceusetesss ses 67
MELAS QnA eli eacenrceinneedeee ees eo 97
Metasesarma
TART Led Wiemrecreccor Pecrerexceccre Cre ree eat oT 70
ROUSSE GUAM ai secehtstess seeds secossassobsasaisdeesiee.cee- teem te ses 70
NU QU LOS Gyre esi een ew Rataseas dicdeleedenncasieecuseiceveeen oases 70
Metopograpsus
TESS OF, vavovasscchac icskscuiies tee eadcs ies ant oie eee 67
LRU UR OI eosecswosccs osvectvew te tistre cic doc RRR 68
Micippa
TRAV CANNON Aes csasiivics Peters eT 27
GUC OMereree tare cccascs cepeasas su snoven sass nsessccwnsesceesececee ce 27
108
Micippoides Angustifrons ........sscccsecsseessesceseesesenees 27
Micropagurus polynesiensis .......0.0sceccserecsseeee 18; 78
Militaris, PetrolistheS .......ccccccccssscccsscccesseseesscenees 21;77
TIUIMELIC GH NUP SIG) Jeaivecsesscesonosssensssovsssucoos snseesteues 26; 78
MINIALG TAP CT Gi svscise ssesoncsnon tence scnsersrcesnegueaveecgaes 59
MINUSCUTA, TRALAMIIG ....0.....00cccercseesscecseeseees 35; 77, 78
minutus
GCOIGINUS PRS socrsons i Mooseseseieenenc 6; 15; 77
PGCHY RT APSUS)...:.ncicencsxs.csetieerglnat hess 68
Mitra, POCELANGA ......ccccccccessesseceeesneesensssneceseenseees 22; 77
MitslenstS, ThAlAMUtA .............cscseeeseccccccecesessseceees 35; 76
Miyake, SAAAYOSHIA ......6..1..ccsccescsscssensesessnsessensnsenees 20
MOlONG NGO 0) ccsacccsscocscosscesssesesnescsnsenseuessensgsasaraiet 97
MONOCEFOS, MENAethiu .........00.ceesecceersceeessessecessconees 27
monolifera, Porcellana ..........cc1scseseceeceersene 22; 77; 78
MONLICUIOSA, LIOMETGA .......1...00cccvsscesecssseessseeseseesaseees 39
monticulosus
Cr PI Od CS irra oo ia ansneseanenssasnnss omecbsabratehns 39; 40
COR OAS Pe. vcssscessuesscnsssntovsvslooevsem esa sane hes 55
AY MODUS Ciseescattscsccutiespeee Maczevececoerseenntlecs 3; 55
Mortenseni, DEMANIA,..........0cceccceeseceeseeseceececascsscennerss 98
Munida
CINE C OR os oss ios sk sain siise eee saie oe 96
CU COUSS OI -osbiaassieke. A saiecsurrteen Ga Nh. caer icoras 96
EV OTOP Mi aoe ches voscscalenbce eM eet ieee 96
iy SUING ois cate oes ect ace ascasesseetesssiasssastsceasdurdecaues 96
VCTLEGUT ONS ee es sei acoc cack souss eomee eee 96
VONGICNELOS ose vcccucesssnsevisesestssnvittnetecotionManscrs elem 96
TLOVIMLGM Wisse ce seen chs sass ease ee cea a ee eee 96
CICCAN TZ SR UE GA NT Se heat 96
DASUREGY es cocsccaciusesascosst coset snnerwaaee tence aucee nde te 97
PUL OSIMANUS occ cccensicccssesessssesesas sates toneseeencece yoda 97
PV OX GUT Apes sal sid ae sect sensi ssccssctessesscuatessteietaneecaae ee 97
DOW RO Cppsreroisssietrattar iscsncsccsnnodsusasuins denaasnscaipebaagss 97
PROPUNAG -oecvsccscicccsectveies cess sisstace sevascsentteeet ties 97
DUST CII Gh ist,« sus s5s c'ssnasascncsinon = ta eopa dts wae shee 97
GUDOUG on. taccse sven dee csaucn ies Gesouenssav ss wee oe 97
WUD OV AEG Bern seas Aesdhncescs ssncies ssecsvinssuscteesscoreeesses 97
SOMO. cis osaacccvexsvondeneusesucsteesae ds soso a seen etind ees 97
MUrOtOeENSIS, HYPSOPHIYS ..1.....0c.ceccvsvrerssversersesseseees 97
MUrSia HAWGICNSIS .0.....0ccccceesccesscensescessecensscessseceaneees 97
MUFULOA, EPIStOCAVEA .......seeveeesecerserereesesesessenssessenees 98
N
Nanocassiope ODIONQA ........ccsccccesesscsessessssscnssnees 95; 98
Naxioides
PCCD sé. ccasssvecponacaivouavencosnedsanedecsiacesseateak eeAeaeeN Ines 98
VGILGHIU, J cacsvaccasnasesnasossssuncesnncstttr aerate eae 98
NeCtOQrAPSUS POLILUS ......cceesesessecsessesscnscseseseecaseneeeses 66
Neoliomera
OMIGIL ccs sssensusvunsdasaavonss vesoces cases Sea eeptnen eacees 40; 41
PPISULGTAS sasiessancincssosscoseeeuevit sede een oe EE 40; 77
DUDOSCONS aasisnsevuscsscevosstusscststiecvescieie ees 40; 41; 77
TIGTHL GUSH... cess shes saccaasteeacicessassa etaasncols eeeeoaaeartot 4]
SPs ssncw vues eosesennectosnssass socinnes dasccete tens ce eee eRORSits 99
VOM OLOS Divas ccsevascevvensveseesaveccventecssincc Rees 41; 45
Neopetrolisthes
PRACUICLUS ac; cicsucansacunsuccnsevuscnss<ensenrsenisteeeeen 20; 77
OPS UV ovata cadsebeves vescvats vacousavesstenecescunnsscieene 20
Neothalaminella, Thalamita .......ccccccccsceecsseeeerseeeseees 36
Neothalamita, Thalamit]a .......ccc1ccccccccecseeseees 33; 35; 36
NGO XAnthOPS CGV GUUS: tcc sviewazsissviiossisazeiarresteareees 50
Neptunus
lOAGISPINOSUS x. sctcuvcd sass sssehcisssscsestssvetsecsrorseeseaes 32
DOABIOUS! wecusiwssscvigeversiieisavisevtnsevinsseosesersoeee mae 32
SOND INCI Si welevnss 01 son sient asuecsspsuseanoreebeneade 32
WIQEN, CHICK OUWUS o-csicerarcinssseussosieossonosioctommpaniameite 54
Migrd, ChlorOdiella .........cccccscssssscseesesseseseetecseceesecneess 54
nigrifrons, Tetralia, Tetraloides ...........c.ccssccsseeeceeees 58
NIPPONENSIS, POFtUNUS .........1cssessereeeseesserseseeeee 32; 98
nitidulus
IRR YM OAIUS ce Oeil coovnostvtnapeaenetees Sh)
PUL OCIS MPO. ccc ceccsezecentesscivass ieveesauneestes eee bp)
KAN AS EON cv asessscavcsasscnsustosevens cates POS 7i/
MANN OD CS). oac. ccsccstvccve assticssonatbcasi cesses ee 50
nitidus
CGICINUS rake heckies sease ik sO 4; 15; 78
COLOPITUS PONE. iii cteoccr at aseiesieses ee eee 29
noelensis
MCU GOUSH Ras. cccccvctisusntacctues sevens sone eee 41; 49
PGF INCA GEUS si esses cessvasessssssiossvescessuscatcuenteareaeeeee 41
MOrMANI, MUNI ........ceccecesssscsseecssenceecnececeeceesceecaeess 96
notatus, Paraxanthias, Xanthias, Xanthodes............ 50
NOLOPOIE!S Tats .......2.20..c02scor.o<once0cuetovssoes br eecuneaceeee 97
Notosceles
CRIMITOTIIS ROR hoc cvsisa sate cnseesase eee 24; 97
VEGA Cr See ee ania susessisdescsesots sac osseeececheaeeeeeee 97
Nucia
DONG sess ccscasssevscsivinsscouSstiace teaveeeseteses eae eee 26
ERY a nd Renee COE RECO eee 26; 78
nudipes, Macromedaeus, Xanth0...............0100+-0-00-00+ 50
NUSSIG MIMELICE .....ccccccececeseseceeeseeeceececeesesceseeceees 26; 78
O
Obe SUM, CArdiSOMG .......csscccessececcsececesssecesesseesesaeeeeees 65
ODESUSN A CLUS ssicvssssesssseesesecesese0ceccssasssczeaeverueianeeee 61
Oblonga, NANOCASSIOPE ...........1..seeceeseeceeseesceseeeee 95; 98
obtusirostris, Simocarcinus, Trigonothir.............00+- 28
OCellata, MUNida......ccccccceccesccsecsseessessseesessecssseeseeseees 96
octodentatus, LOPHOZOZYMUS .......1..s..0c1eeseecereeseeeeneeee 46
Ocypode
COV ALOPHAhAlMa .....c.cccsecesseresseescsesseeeseeeneeesenenace 73
COP AIAN RA Bid cssciasteacdenaenn eee 73
LGOVIS ccsecveestius vansastan saee avons tateesicate rescence 73
MRACTOCET Gv sce coasccoeisvnnvanse sues tetee cee OS 76
Pallidita er Oe... wescsccnssssiccencecehamieaes 3° 3
PIAIEP SISTA scicoosssssesdriees rn eureee 76
ANd (4 wean eee Ree errr co: 73
OCYTOCWA TET BALIS ono snconacsvancasntacnannnesenneensn tute 45
Odhner is TWeediela icccisiininanincinmenaremmer 7
Ohshimai, Neopetrolisthes .....ccccccccsesseseesseeseeseeseeeens 20
OLVIEFUKCOCROBII A skxcccecdsceussscesuxcantsucreacuentess I2Z<13° FF
OFBIGUIARIS\ LASS OGGVGIMILS acccccutuvsccieest eons (eee ceenetnCReyS 30
OMDUOSIRUSS POTUUTIUS \.snacesccscescastecoesecvatecersceeuces 32; 76
Oreotlos
CACY MUS Riscssicsisiorenesaonsrtimmiggouretaaeemnan 97
POLARIS RAR sisi cctsorincesst ici coer 7; 98
orientalis
Og ee 31
Homo lassccsescasceceei Risse CRORES 97
L@<—ax ann. eu
ornatus
Gy clodittsien scscseheatescsttesese as OU 55
PR ONULU US ticles etaede feseeledoett Sie eee nce 10; 76
OSaGHIT aN ie casns been Minis Sisica\S oo eee ee Lee 28
OVALS: LI PPG snvstet scar titet sociated cee tau ee 5; 23
Ozius
PAW AIENS US eeeeereere ten Vaca an) cleus ELE ARS 64
HODGLUS Baas wank relished lowes) 2 Morin he ORR 65
TUR ILOS US ist rots ree siasrs inked shst earn tA naa? aire 64
ETL CAT UIUS ene innien Secs see A Lies eee ee 64; 78
LEU CALLUS RRS Pa CRRA ECan SL er 65, 77
LUDENCUIOSUSR Ee EARN aia e eetan ae 65
P
Pachycheles
ISOLA Seer ot tee stately sot cestinscstenctee em cee eee 20
SCULPLUSS. Bhs sco leah ioe icra ed lace Wn alee meee 20
Pachygrapsus
PAR ALAVENSIS CLUE LATE OSEAN HIN 4:68
(OYA SERY OER te nee Het eee eRe Pe eo EM 68
THLUTIUTUS seein rasecsttee ane tecees nr tereon cater eet neac ree 68
DIGHYTONS Borer rets feces fe csuie es ese anes eerie ose 68
DUICALUS Eee ri a renee eeu ue eee es 68; 81
SPM sic cese ess eccureviccrescasr asia: easenssi teeaee aretcu ones 99
ASV CT SUS ree tN de wencest ctr ssacestice eter 76
BAGH ICANT Payee NAR N or ee ene en ee 23
pacificus
TL GUIDS MATIUS sn ssass esses atest iaivas esac cect eee 96
GQuOPG er Us Aer Nes. aula etek enn een eee 96
TE AHODEUSI Ee ete test tereN oe writes teste err aeeeere 51; 78
REPOS Mar eters ees enna 23
Pagurixus
GHIGEDS\ Macernesse enc coeen ieee eSeces eee geet cietan sete teen 19
LAGVIMANUS etre ene aren ett Aen ea Ee 19
LLALOL OLAS Hh SAREE ST ate Rae Pen Re 19
Pagurus
GUI CHIUS Te recs trscced truce sce nite en tne Rect eraeae 14
EONS rorya ereeoe re eee 17
CNT OAL TIO reed nrer ner eter ee ror rca a ee ene Re 17
COD SIS ee eee te nee NNR ee Rea eee 18
DEMINGALUS Sn eM ont ene aE SM es ce tareetver esse tte 17
STL CHU AS BARS SORE CH eet BEE REE REP RES it eA 17
LYE LISS Soiee Raton een ree Rao athe 15
DUNCIGLUS Mrrentrnets atte Mercere erro: 18
SONGUINOLENLUS ise nial hes ctk) abc) cr eee tes 18
SPLUPUITLONUS re cast oe eee Oe LN eee seen, 18
SEREO CLUS rina OU NY ER MERU AUNT URN BANAT ERECT, RENE 16
Palapedia
TIGR QUCSES pest ci esen cn Sica crest ontes Sotvcesvseaeiieeiss J; 51
RASUTIPCS MO crete ete en her ean NY MN nse as 51
PGND Yy(RUSMARNIFICUS oon e ree cesses 7; 10; 96
Palinurellus wieneckii .....c.ccccccccccessnseeseeseeeee 10; 76; 77
EZQUNMUFUSNSPINOSUStee ee eee 9
EGVIAUSTUS UNICONULUS arte nee een ce 96
DGUPGARETOMET GERRI Tes eee Hee 39
BGAN OGY pode nese eet eS 3; 73
pallidusi@arpilodesimrercn screen 39
TGNODCUSIDACITICUS® Mot wet ot SN sheet 51; 78
Panulirus
NGSCIALUS Meese cre tn ae a otneis ee os 10
109
ROMGHUS siececsecscdstesccsieticcssah sis Pn 9
lOngipesiiied Wee decay Mone eile ee ela 9
ONTIGLUS RR De eee ce aN cays 10; 76
IPASCUENSIS eh aretha essed laviaiesse eee 5; 9; 81
IDCNIGUIIGLUS RE rere cen ie Ue eee an 9- 81
DOW PRAGUS HE sree tees ee 9; 10; 76
SP IMOSUS ire ennsnsceale bettie sta ANNO ST ANA 9
VEFSiCOlOT asiuaie kes Wessnerlate AA MOAN en TS Le ules 10
Paractaea
OXCENIT ICD ee cacnsUnasedisestusactesviinco te 43
GUGAiIGrCOlAaL ane ee eee 44
TELUS] strane bene nt Ah Lu lek acto ci MN UR 3
FUSOPUNGLAL A Renaesitesri i inieiatouberd eaniaans Rae 44
TumUlOSaMA veers abieliy aerate Noel gates LI, 44
Paractaeopsis
QUAM IAF COLALUS .....eccecessesseesessesecsscsssscescessensasnaes 44
CUM OS US detehesdias ei heatesaiveviee hicdetiaes Rares ee 44
Paramedaeus
TLOCTONISES PEEVE ON RRNRI HUee Lsere ae st) MEA 41
ISI Lexibase tals usesenrsiah tes donnie bots CANDO 42;77
ISDN 6 ORIN MARU Gok sD, whey xaSatees tata tiaes MAU EARN 99
Parapleurophrycoides roseus ........0-. 73; 77; 78
Parathranites hexagonum ......eccccsccsccssessescsessssesesesees 98
PAT AXANtHIAS NOLAIUS ......ceccseesesecscsseseesesesseasesessenenees 50
Paraxanthodes polynesiensis .........1cccsssseseseseceeseees 98
PAhCQhMICippan eet weak ee 27
Parribacus
APU GI.CLIGUS Wee he Need one te ne Nee SAN 10; 11
OURS PAE IE OE eee eleueeece 5; 11; 78
SCANLALNUS EINE TMs waeaooneveeasea eee ee 11
UPSUS-MGJOR As ee hens Il
Parthenope
CONINGTIUS cssernite taeeeed eel ace ae oneseaiteg AE 28
Hoploniotus vous. terse te pe ccins tense talks lee ee 28
hornidans sel eves eee, hovsatecess tet hhc nie) SAUNAS OD 28
LAMe lI GCra Ren rr Races cnet tae sac 28
IDELA SIC USC RN UN Rep edees antl clahetn ecloa ko Ma 28
DOU IN ieee riage. Noe nil rortea NT 98
ASP ELS Mebe yo rated span uss rela ieee earidtes BIO MERMAD ), 99
STE LI GUGM meshes oiha Cees cA ok 98
parvirmanus, Macrophthalmuss ........cc0cccccscrsessseseeseses 72
parvula, Actaed, BANAIEIA.........cccceccssessescenseneeseeneees 38
parvulus
Cy Clograpsusrcwesauncdeleecetresinicoees 70
Pilbmnusteiete vetlel hl cit, NS MO, 62
PASCUENSIS, PANUTINUS .......ccceccecceseesccsseeteeeseeees 5; 9; 81
BASIUNC GMMR IAG Renken iene Sie eran cats ine Aa 97
paucidentata, CHArybdis ....cccccccccceccessesceessees 7; 31; 98
Paumotensis, PilOMIUS..........cceecceeeseseeneeeeteneeteneeenees 56
pedunculatus, DardanuS ........cccccceccsscssssecnsssseeceneeees 18
pelagicus
ING PLUMS Pe Ri we we ee narnia ak AT De 32
Rariheno peri iie sak iihiconri teen sures lass eM 28
ROT EUTIUS Ree eta ener Leeda ean S277
penicillatus, PANULirus .........cccccesccesscescessecceesecnsees 9; 81
pentagonus, Echinoecus, Eumedonus .............000000+- 29
Percnon
ADDTEVI GLUT ee sehen a entencn aucune SORE aa 71
CT Reaktor rr EL CHEE eerie on ee 71
A Wiechednorcte ectte Cee ce EE Ee ES 71
PUI OLE Sr enimch slr uncu shit besos BANA OE) oe 76
Diliman uspiscen ene hove Wnts cectenics oben Oe AE 71
(PLAMISSIM Um ett serra eeceetsaentig atria culr cian EEN ae 71
DNGRUSSEMUSTO RES cc OEM Tass cccsecnengececboas 71
coe NE Ee ie een ee a en ee ee ee ee ee
110
PCLVOT PSUS CRCEISUS UA, fcccsssxcsewcses sven cwesosss iaaanansnaaee 65
Rérinéa Wimida Yn aeccintncdnnawe ees 27
perlata
COCNODUD . céssevcicctescccccessuesdvotes eesswwsapengeatae 13
DAI QI vcsccstcuvatisslstitoceteceveceine sc mgeyee ee 75
perlatus, COCNODINA .......eseseeteceeeseeeeeees 12; 13; 77; 81
perpusillus, PlatyOZiUs ......1..1.cceececsseeserceeees 75, 77; 78
personata, HypSOpPHryS ........1.scescsseccreseesecseenceseneneees 97
Petalomera WiISONL .......sccccccecceccsstsesssnsenseesecseceecnaeeaes 24
Petrolisthes
DISD INOSUSB IRON iacsecisavecccouresitercsas ec cun eo ertipe 21
DOT GA QUEUE wvvsciesneacsbes sre savedveves Caees tees 21; 22
DOSCH RS sec cs ei Nec sav aaces tie scstbc cneacstauaaiangets cas uc 21
COCCINCUS IAD FARE clases dexesenesustexecscenien teed All
CCACENTNUS OOP cs caccsteeeneonieatees 21
CLA ed GER OI cecrsccteuevenss corseyreeetehornrmemieee 21
CLOG ONS oo ca vss scvecesssesersttisnsssicsuess sient cee etaas 21
GOAT. GRAN seessess os secewevecsavateitenestesie tale eave oe 21
ETT TIS Sern OOO oer 21;77
DUDOSCONS 0. caz scsssusoss sxicurevensassvavsacasisdesoso eR wee 22
Uf CSCONSEIE IID svcsvcvvestensansseeceiceuusensvensens PO Ae Tif
SCODRICHIUS:. ot. ..sc0cssssceeuoasssucomabsnieepe Rea. ocah eee 22
SPOS, sean iseteivsiaesesvorwssacrenurermaeetabeest hae aein es 99
LOMENIOSUS sce cetcessssisees sees enhernteinces eased 22
Philippinensis, Thalamta .........c.cscceeresneeseesenees 35; 76
Phylladiorhynchus
DUSTUUSIS cosseciscscsseessccsvssvsveisesaeare errno mate 20
SEPT UT OSUPIS secctes tees cece satennssuritiaayseniev Recbwa regener 20
Phymodius
OT ANUIALUS sacs casetevenesansdcunixrnveveranremnbumet 55
PT ANUIOSUS ica. tersiicnseurtaris Aves etree epee 54
TLOTUIGT COIN ere Ene Ore 3,55
LATELY Wieereenecemtcrccerrececereteerrcrro erento carrer ened 55
IS Oe ecto ees tt veai cape den elceule Hor arsersveesiewenemeuaralere 99
UN GUL GUUS cosets nso a tren evieitet ieee 55
picta
ASTON Cl sieccciosseecuswcc shove. susvecn sé stceir eee ete 25
IM GE csccses ta viee tian ioubaset teanetzenveeenerstte cs ietaaen 25
GT LHe atone oe racer cea oe eee nes 35
PICLOFILOPROZOZYINUS F.ncc--2ccssccaseeveceneraccnn-oeopets ee 46; 77
PICS SGP APSUSK vucvievcves leptesceh con road Moran von easy encuck 67
ULI GNU SOP CF CRON ae ccccecnsseeestas Sar ee ant ee 71
Pilodius
QDOFTGNS ev ccncosccsansaicseresossncnenssinsust eran tage 54
GhCOIAUS sc swecsssservieeseernstrisieen sone Meare 5)p)
JLAVUS isso sevicni ooosnsever0ssessseas<ooaaepeenaneep Nts anmereeancy 56
TRONS Uv scek seca vak cnc Geechee wave sec toct st eee RS eee 54
PAULI ULUS MH PEE. «sna: 2 sour a Spahis es eee wae RAR 9)
PQUIMOLENSIS ...5c0scsscccnsuseresesi023 Gashbcrat NM ces tape dts 56
JPUDESCENS 0 vcosecasicsncescsncasssianyebs¥unas MuenousteRicsccses 56
DUQUE isccnascsesnsesacvastancesesecacvcusassenescaestee rates 56; 64
SCADEICULUS EES cctaciscustec consis ce es 55; 56
SIDS sinicoosceatsssececessts vactsssecevensecsscvusteepenca tan eee 99
SPDIUD OS. vsessssassszonsssscasnnnsevesese nes cstsontacesmecass eee 56
DPUOSIMARUS, MUNIG. 4. .ivcccconconiecpspetteencec hese PO
pilurmmoides, Thala itd). We vescccxcsssk- ee 36
Pilumnus
IRLODOSUS ccessnassicsnsasdséseoissisncecie tens: «scene bser 56; 64
LOR DICOINIS .sscccocvsxesnduescorcovidenc cesceusesactenio eee 61
DIGI DAN UGLUS 0.600. ss sesinsctnscecssssicsGson cork gaceincavtag 54; 64
DICTOUCNIGIUS, 0x ssarevsssakvasnanssvasecsesssscerseao ence 61
DOTV UU S: sosevcncencscecskencckesannicanccangacccente ean 62
INO Lot Rrotc ot ee 5; 62
Sf) DN, A ISSIR a vssususcodesnceseoxcusaae conte De seas 99
LCITI CTS IS OREN ooaiole soca cal eedeas suisse ese ee 62
Mpirinotner eligi |ACvi Ct eee cessecan eo nsnoses eee 72
PISOIDES NP ACH Y CEI OS aon oon ace a nen nnenne semen ee 20
Plagusia
EDN CSSA ove seh sede eoehs fussed neste ep Re 72
SULIT NTT Rene AY Ree Ree, Bel pce 72; 76
SPECIOSA PIE on scto ote os secary tee oepsstesatoanl cae 372
SOMGINOS Gre: Aas ot RO dso cssss sve ee 72
BUDCT CUI ON OD ioitees sac ones scstcscanscdeonssoaneeiastucas oe Ee 72
NGOS CY QNEUS \o.csa0.c3.cssneissdacassescveossasvaseeesasapecaae 68; 76
Planifrons, PACHYQrapSUs ....1..1...cesceeesseereecersenseeaeeee 68
Planimanus, SYMPAQUrUS ..........1-eceeecsecceeecnecneeececseeee 96
DIGNISSIM) PEN GNOM aor stess-csce-seeecenszeoesesce-toee oe 71
planissimus, Acanthopus, PerCnon......1..c.10000000000000 71
DIGNUS | PSCUAOTZIUS sascsctsaccseceessteescoees oe ooecee ee 65
Platepistomma DaISSiW . <.<<..20-+-ceneosossesoseeseaanene-teee see 98
Platylambrus, Parthenope .......2:ccccesececesseesecenseeneens 98
Platyozius perpusillus ........:01.1ccscsceecesseeees IS TI TO
Platypodia
ANG GLY DIAM. 03 sss cciseseisas'snsvascievecsuiscetaessinagtuasn cece 46
YUNA DS RS cs sossecsezoenensxstontnsetentes ane een 61
granulosarihres:, 2 eal hi we eae eee ee 47
|PSCUAORTGNUIOSG «2.c002.secsscen2nsassnnepecpeeeguere eee 47
ISCINE BY QNOSG xc as casts vas socececouveseotsuoccs ioaeecz eee 47
PlabytansisOCYPOde o..2..c2o01:<0-to2sonesseceoenneeeeangeeeeee 76
PLESSIS CHOP ASUPUS, <ccceseacesssecsaseusessanccaset ghee 96
leurocolpus DOUCGUI <0..2c.2.-<c0c-soesosnsvoasaie eater 95; 98
DICKGUIG, MUNIE Gs crccscexscocccecesonsesose-¥exs-ceeeg ee 97
DUGQIMISHE GCI OT ADSUSp cereascoeee toca eae eae 68; 81
plumosa
DDI BOW sein ssc coccaeis covevenstavecetsvieecasas eee ee 37
PGT AONDOG cs. ccnesensvetesiescassnasecosdsseussVase-aa cee 44
Rodophthalmus) Vigil <-...c.c.x0.c-s0asersccscservass-<eeree eee 37
politus
PIS TODSUS sx. .tcesctcsie cess oovusxeversaxeseucetentnees 4; 66; 70
IN€CLOSIADSUS: soca: icccces3 want svos<ivarcocsnttermeneen ae 66
POW GCONENGVACIGCE 120.00 :0e00ceneecerconceannnss 02a; epsseaete 42
Polydectus
CU PUA lifei asso 25 ssn s oneosvoresezcoitienesernnetepae eae 37
WELL OSUS Fr aerscscscbsrrsescet es 0c Sts sS eT fee Shay nose yaane 87
polynesiensis
NY HG Ro) UST A Reece a ee 18; 78
PON OX GMEROUES . .-..-2< c.c<05s0sacss3ca8<0sscesuees eae 98
POLY NOCAMURNA GR... o.cc.scesscsecns csstasosennssrenseersau eee 97
POLYPRGBUSHPGNRUTITUS x. ...:.s2n<cccancsssacceussnsncossupentel 9; 76
ponapensis, Lachnopodus, Paraxanthias, Xanthias. 48
Porcellana
GCOCGIIE Oe oncom evo nsunckntsh aonec ana ea ate 21
DIRE LR Lda A haat scare Shee L297
HOT ON J CNG renss vcr cavuscxsecuetes certs. coesvse ar 22: Jiz@e
FV OSCEIST 6 o.oo ccvccnss0k caesnansGnensskcaneeceeenesegeetne nen 22
SPOCIOSGER CA cs .csn-scgseceh cctnesteunccenss sc sSeae see 21
LOMETITOS GaSe Si REE vans scasexanaseatecaciny Santeuaaaanialt 22
Portunus
OSA TU ROPERS ao as wane canteen ee 31° 7/394
ST LTTE a cc eee ee ME - 3]
17111 (21 [ere Ree ene ee 3]
TT 112) (4 (eee ere eee Renee Seem eee 4; 3]
DUO io oo cecen an oucnaneniar ghana une Rca SIZ
DIED ISPINLOSUS soos sox ccsbacssnyensees cect anna MERenene ee neEaES 32
AECTODIERGIDIUS so, Sonxsnansseasssycennssestecntentaa on 32; 76
ID DIONENSIS soc ccxcivacst seotics-sheene caste sasaen eas aes 32; 98
OF DULOSITAUS wcesssicckanssuceeuescacesetn teeta eee 32; 76
OO Oe
DEIASICUS RE REE OR BN N iecossashieatuees 8 U7
SANOUIMOLCTUUS ret vencsen ee eira neti h scene 32
Potamon hydrodromu ..........sscssceecceeesecesenneneeneeneese 77
IDOLANUSHONCOLOS rt cset ese eek cee ence 7, 98
poupini
(CRGCCONM eho core aec sa Nak oes hes boas eiaea saree 98
Par thenOpe ve. cccecssesvsstas ites sas catctin cs teectesoateetesteas 98
SUZ OPAQUTUS =sscc cio cse tee Svsesetiakeveaetaeicee ese aea 96
SIP AQ UPUS od seteke erste niuccecacscassetettls acne eRe rete abe 96
ROUPINIGIAL SULA cert ci ca cosesosiesccsesoces cxnsstaneionaatisctassetice 97
Praedator, DYNOMENE .........ccccvcsesetessenenseteeteneeteeaes 24
JOY OLLCLLED NG TED oscar cece COACH ECC 97
BY OQEFY ON NAIALAE 0. ceccaccevsisesiesovvinsstunnocusenasectseigs 98
PROLCUS STL UON Arete tierssescstro arecace aa ait ORR ae as 27
Pinyin wala laa ee een eee 36
Psaumis
GOVIDES tecorcets csc sticrcsetsek een shesdestentvuseelgseahesie users 44
COLMIOSAT EE. erect sts cortsciesieshaskeisicean tao aaderagee este 44
pseudogranulosa, PlatypOdid............1..0:c11ee1eeviee 47
Pseudograpsus ALBUS .......1...cccescerenceeseeeeneeeecnetnsenees 69
Pseudoliomera
SHANOSNINGNA aecere tis ecco devo nsiedases de aceoteeecedses 44
LAL REE RSs ORR AMO L SRST Ne uegiees SePar vente Tet 44
UPB CLIOId eS meee ese ne eect eter eee 42; 45
SPC CLOSG votes save esrsepan seek cages O2ie Ue ceas ean tie waete sata 45
WEALD (CT eer nec ere eee Oe rte ES 45
Pseudothalamitopsis, Thalamit] ........1ccsccc1eere 33; 36
Pseudozius
COY SURUS NS ite tetera RESUME e unset irae ue got 65
JOLUCEDY Dhebncrpreerctse heorpo ee eo onc reece ee EO 65
Ptychognathus
CRASSUNGNUS Ee eR oe eed 69; 77; 78
COSLCL QNUS On pre Ore ee ee 6; 69
TRECH INC UIUS Ie nearer a nce eater eh estan 69
pubescens
INCONOMERG e hrsWer see pelieeici aris mahie, thes 40; 41; 77
PEL OLISth eS iis srsrscsistsen tices ca scenc dated ctor eee gta 22
PVLOU EUS ret Or mre ae ante cs Fe ERS Pee Teac, 56
pugil
ChlonOdOp sisisasnseteos eet ee eis eee 56
TRULOCUUES TR rn OPTED Ae ORD Dh) SEIS Dealer 56; 64
Pulchellus, GElASIMUS ......ccccccsescsesesesscesssenesesssensesenses 74
PUICHRGNMIUNIG rer rere eee ee 97
PUMLUSNLOZNINOUCS retraite eee nee ee fate 47
Punctata, ACLACOMOTPH| ..........sceseeeeceeenceeeeneeeeeesens 28
punctatus
EEISUS rr st eee ea re ee ene cre 53; 77; 78
DOTTY UTES ofa ARE esc Ee ea rt SI
PUNCLIMANUS, TVAPEZIA......1.ccccsceesersecsesseessesees 6; 58; 60
JPUNCUUTALUS IP ABUTUS cc cccrcersncsteseccesest ssterseteotostececenes 18
pusillus, Phylladiorhynchus .........ccccscecssscsessesssessees 20
Q
Quadrella
CYT ENIAC errr er re errr renee 57
LCWITISONI Reenter eee eet eee 57
TACUIOSA aE ERD osccreiei be crttoesiteescete 57; 76
SPI e neem eeeare erates eee eee eee sen cdot an cevacbuaetorennseiects SY
111
quadriareolata, Paractaea, Paractaeop3Sis .............. 44
quadnidensMihalamit apm mec ee eee eee 36
GQuadiilobalawiGlamil arenes eeese tea DO
GUAT IIODALUSH ON MO wareny ee tetetens eee ee 38
quinquedentatus, LUPOCYCUUS .........0csceseeeseceecesseeseess 3]
R
IRONIC AUIGISERENE Neer es orerae rene Se 98
1 ROTEIB OT) (ACL 07 Iobcectcor ence eReeOct CEC PERLE HORE EOCEIoa Hocecen oe 24
ransoni
GUIDA AI AUS irs ean oace eee ee een a 5; 16
] UDA TATA ee sceoubecesceeo eee OO OER eSeop 5; 62
TRAD ANUSPS GY LG US ence tece ed eannune eee 95; 96
rastripes, Kraussia, Palapedi..............1..s.scssceeeeeeenes Sl
IRGLAMUAIMOLENSES Fecasccsseccasssecanu cs Soo ctes feiaeas sot Ue 98
TED QUSWANV. CUES eo were se cence acdsee sees sssess eee 10; 76; 77
Remipes
DGGUICUS ES iis soos ecieiai) iin seo ctseeueye eR Ae oases 23
RESTILCIN AMIS pc cessceensssssscdsscarec cassette 23
MEL GUIALD Ny GDOZIA ey eee tee ee eee eee 58
retusa
CGIMPOSCI ooo ci ccees sieis eicvacstsceh ve der diewcet Lee 26
PGT GGLAC i siiat vaceccusseronseses ho aieeam eee sedate 43
rhabdodactylus, C1UBANAPIUS .......1....c.1cceescrnenteeeecenees 16
Rhinolambrus, Parthenope ...........c1ccsscvecrsvesseseessene 28
richtersi, Actaeodes, Liomera, Neoliomera.............. 4]
rimatard, AldinNOdQeUs ..........11cs0ccceceeeeeees 7; 41; 78; 98
TOSCORINUGCIG) Sah OS ec vac Sos vou eect aahesscaientasaeoeens 26; 78
roseus, Parapleurophrycoides ..................++- Wo VS Ths)
rotundatum, LADUGNIUM .............00ccecseeeeeeeeeseeseeee 70; 76
TOUUNANTONS) CAP yi Gieecacccecees-cnestesesceese eeteeeeee eee 29
VOLUNAUM, CAL dISOMA......11..cescccesesccsssscecesscsesssccssnsess 66
rousseauxi, Metasesarmd, S€SArMA...........0s0000e0e0eee 70
GUD CLI GS MUNI deere on.) pecttnsoene eee on erect eete 97
ATV OE AOE | EYOTA NG Rep SSO RO ee 39
RUDOVAIGIM UIA sc. -sscecsccssscessekessces soto see 97
yueppellinGaillardiellusyrc cee ceecesseencsesttes 43;77
rufescens
IRELTOLISt iE Simetnihiitese noni os Nn ae PLS WAST
OSA NATO 5 coccenoro eee OR ERE ECE eee 22
rufomacilatay Mer told) .cvc.c.cc<.c000s.gesess1ceetiestaeensetees 98
rufopunctata
(A CLC rata So eit A core AS SOR can sets 43; 44
I XCTRAICERTT AT Testa eee EE ee ee ere 43; 44
RGD OZIGE Swern stacks tr esseatses coscriassarseeeiet 59; 60; 61
HUES QUAL OIE, Give tice ec ivoessesees ive suenevidedsoeviogeenteecs 38; 39
RUG ALS GAN DUOM CS) res tensor oc cocessn: «sacneedentewtensss 38; 40
rugipes, Carpiloxanthus, LioMerd..........01..1..00+00000 38
TUG OSG CONODUG or. ca ccn sacs ccesenstevescs snes 0aganesaeiieaes «eth 13
rugosus
COEHOD I Geir, cosets sveedicesstet aba cecenscciestRMeee eae 13
IEE LESU Saris es Soaks ooecss tusbicosd eins alacaautendeac ume e 52
NPR OPUS s<o eee sano oshtsg tac evasevcseepouteceueas bvatuceeds 4]
TUSUIOSAN MCLASCSANING ince. scs0510-t.cevsoserseseaonseis sstaeteen 70
HUD ULOSUSHOZUUS, .o.ccz,snscxcivenntostecpolesnes ene ttn sper ahecenetes 64
Ruppellia
CIPI DES woh cin cnssscassaiacs see tinganans Beco ee 64
OV ONULOS Be vv canes anccnneyacusocesiancnnntesteenestn 64; 78
ruppellioides, Actaea, Pseudoliomera................. 42; 45
EE ee ee
112
S
Sadayoshia
CAWGASIR oe Oe 97
TUV AKL cocscs tis cccestiivdess ORS 20
sanguineus, Chlorodius, Leptodius, Xantho............. 49
sanguinolentus
DAR AGNUS, Siren Riera cton eae ete 18
INGDEUMAUS sn o52. sos socescescsctsnscutbscscvesttsisveseestaseessebasiee 32
PG QUIS eee reali tee aaah gen rade Nae eat es 18
PONE OR ae es 32
SOrMALVUMNChASSUM PE ee ne ee 70
scabra
AICLAC UM rn cite RE Et ON Me 43
OL ESAS BRR RR ALS 43,77
scabricula
Ghlorodopsis:iceccs See 5S
DOMeCi a. DE PA 63
scabriculus
Petrolisthes 2 eee 22
POGUES ee Oe Oe 55; 56
SCarlGlinUSWPArniDGCUS. ame Il
Schizophrysasperae nee ee YS HU
SCTUPOSAN ACHR. iio OE 28; 77
SCUIDIVUS NE UNGNINUS 0 cecctees-seccscscee eae eee 4]
sculptussPachycheles sence 20
scutellatus, Dardanus, Pagurus ........cccccceccensereeereene 18
SGvUlG Serrala: asec ee 33; 81
Scyllarus
Gntarcticus nee Se ee eee 10
GUTOR ERR RAR 7; 11; 96
FAD QNUS) 12:52:00 A EIR 95; 96
SD ee ti ed RRO ae Il
Seband) Eriphidiicdccc. 8 OS 63
semigranosa
LIONEL renee 40
Play podia ee 47
Semistniatus) ClDANGriusneeec te 16
SENEK PEMONISUTIQ A): oaz ct eee 9
Seplata il Gpezianarn cate 58; 60; 77
serenei
IMAC OPTUNGIINUS teat csrerccetesescscsttcscaneccs scott 73
Randallia wcities 98
DVFOQD EZ eet Aa a a 6; 59; 60
serrata: Scyllaiwen eee ee ee 33; 81
SEVIQUFONS: Net Quay nce iss cectccseoee essence eee 57
serrirostris, Galathea, Phylladiorhynchus ............4.. 20
Sesarma
ANQUSIU[T ONS Sn ee rae 71; 78
TACQUINOL ee ene ecress 71; 77; 78
FOUSSCQUXL AE Oe UNS tere etic 70
HAD EZOId EA. eect eee eats crtreeee 70
Setifer ACLUMRUS) 252 ee reenter 61
seurati
Caleins:ATe RRR a OTB)
TUGGING 2icctae RRR 4; 36
SIBOZQEN SIV ODOPAQUN Sern trcctcasettereavesseceeeaneteee tenets 96
SLENATUS) AlErOAlOpSIS tictiteerretcrce cette 45
SUNOCANCINUS ODIUSI OSINIS elie cicdcocstetee t eetee 28
SIMPIEX, PAVAMNECUGEUS <...cccecescsncxsesectesrcssatvencettet 42;77
SINENSE; DYNOMENE eiiiiaiss tacdevlezernesee eer 24; 76
SOlA@ MUNI dae Ree 97
SOLiLAriOPAQUIUS SP. NOV. ....scesccescsecsseeserseeneeseerees 95; 96
speciosa
LACIE EID TN Sars ascedceerov ested saepoepssedeottcecoses 45
PENAL CTO) Bea ie pace eS A Ra eee 22; 76; 78
J EAET ATIC (7 tenons anise arene tie ed, eat SI 7:
POxCe Nana erro iis wes nctomoroveseesoacteanernuctus Meas 21
PSCRAONOMERG scrcorssnicsntonswinenssinnwuevacecreeaee 45
PV GPOTi GMa careectecteateconseeocicevasseomereaae Same 58; 60
Sphaerodromia AUCOUSSOL ......1.ssecseseseeseesessererseeseees 97
SPICAIUS CGICINUS .25.S ciceccswcaceccscevecstssetvasecoesteote 4; 15
SPURS C AR roc ccccwactoccnovestecaucoreceocsgadonseee eae 74
Spinifera, TRALAMIUA ..0.....ececceescessecesceecessecesceneeenes 7; 36
spinimanus
RGR URES | SE SL cences soar octeeeeees 18
VGlA Wi NOP SISO, FIN csc ceneoeezewcceee tees 97
spinipes, Chlorodopsis, Pilodius ...........1.ssseseeseeeeee 56
SPiNOSA, DYNOMENE 0.....1.csecssescessesssesessceecnsssseeseesceneess 24
Spinosus
Coma bitia eee ecteeiteeaoes sess 122955 Fe
PQUARTUS AOS echt meee eee 9
POR RUS oie co.ccccocteteenoesoend Deoreictenes nO 9
Splendidis; EtiSus: .2r..i.cc5cc.00ctiseocsevasorsvvsceooeeucevoreeaees 53
SQUAMOSA, PIAQUSIG .........scssesecseesecssensesceeseeseeeseseeseee 72
Stellata, PArthenOpe ..........ccccceseesceerserseeeecesceneeceseenaes 98
SLIMPSONL, LIOMETGA .......ce0ccssceesnoessscconssnccancsnaceasconces 40
SCOFNL, GEOQTAPSUS .acceccesseeeeceseesecenserseesecesseeseceasenees 66
strigatus, Ciliopagurus, Pagurus, Trizopagurus....... 16
StriQOPAQuUrUs POUPINI 0.0.21... secseceecesenserseeseeseeeceseeeces 96
SUFI QOSUS, GFAPSUS ......c.ssescescesceeecescescnseeseesecaeeneceseases 67
Striolatus, CLIDANATLUS ......cccccsccccceeeeessceececessceeceesenens UT
Strobopagurus
BU ACIUD ESS vivre scsn-csocscuscovescess sUosovuecesevereeeseeciees neem 96
SIDO LAER. cs cscsceccocvovousesecuusceavoratencvs covceceurtceeteeeee 96
SUDACULUS, LACHNOPOAUS .........sesseesessesseessersesseeseeseess 48
SUBONDICRTATISN CY ClO oo coc ccecensvcee se cov esecencenot-seeeetaee 26
subquadratus, Chasmagnathu ......1....01ceeeeees 69; 77
superbus
EL OPHOZOZYMUS orc csccccxsccstocees voce vacvervonssceusees 5; 46
MATIN O ios occ cvastuavsntsu ances socduscesceeettneneucessenescee eee 46
superciliaris, Actaea, Gaillardiellus ...........:.1.0000000 43
Sympagurus
APPA ISTO coos ccscouseccesseosesreoureeconcosecussesthestoaae neem 96
BOLLE CTE oon scccawnntsacvesevncceswoncocge chance ceeeeeeee 96
DOUGGINVIN ETE oocciexcoceseeasecsvnsscopsveceee teen 96
WO PRCTTIE REIN ccccoccneateonsonccsomssonepeae- tomtom 96
PI ARITARUS E eo ocesccccoancssncooncesuonteptectcaeaee 96
IPOUDINE, B86 COREE CAE Rhee oeseteccrneenceeetene 96
URESDIROSUS <5 cnc. ccae ionesnvasecevseursses sotuesces eee 96
LUTON NS oasis ies as cakacnecaus satcescte eens emcee eee 96
WALLS ih. Soiicinccowiace scr wenceuctu naar eneepenay Tete Sea 96
T
$AfAE MEP ACTRE GT rracacaiwcccsecontacs vetotatauscemntcnnn te grecees 98
tahitensis
EGchnopodus, Kantho sovsivcscwcuscccreeraseeteeente 48
Pi LRIIAUS sees ccoviccnervesverivesndeseneen teeter eee 62
LANENSISHDYNOMENE co.cc ce ccreweeoeeeesceere ea enone 97
1CGIULP NGXIOIDES cscs cocscscevansecevs cccseteenecenecuepapeatanteas 98
LONIIGPUSTAIUS, GFADSUS%.ccv.ncccznsecxtch erect oats 67
lentilf Ons, ACARINOPUS ccc ven.eucs essence eee ee 71
————— Ee eeeeeeeeeEeEeEeEeEeEeEeEeEeeeeee
LENUIPES CAF UPA i002. scsvssesccaoe sees 29
lerrae-reginde, CAICINUS .........seeseseneecsenceeneeeees 15
tessellata, Ly bid, MeliG.........sceccecscecssecnsesetssnceecseees 37
testudinar ius, REMIPES .........0.ccsssesessssescrsescseeseneeeess 23
tetragonon, Gelasimus, UCQ ........ceeccseseceserseeeeeees 74
Tetralia
COVIMGNG irvccceneracesnsnarorethcs chonstinecvenruiees veneele 57
CINCH ES earl vic crease sstetsiscctunsctstsausestesianesevesses 57
QIADONT IIE wie scovn cits ccovscocenct cay tesesvesessrsscusstcossensesst 37
Helter Od GCL La eee vrescccsscsrctvtivosestestescceasstooseeass 57
TLL QTASTONS sree car conc coecresten coca etek rete creche seen 58
SOFT GLUTONS cscs vccsrseseascssessencstestessveesonsessessecsesevets SW
Tetraloides nigrifOns ........ccccccecssccesscsesseseeneceeneeneees 58
tetraodon
UDOT Gee eee ee ee TA ctseleeert ets SI
VUNTAXANINIAS Surette cette cateeseecs eeties hceseawect ay
XN IAS ee See een een 51; 77
MOM OV: tes Siew ieliact uve te suausauouiele cioveusrssdaaxteertr oe 51
Thalaminella, Thalamita .............c1cscsceccerseereeeeee 33; 34
Thalamita
UCT LY CARRERE ee REPEC Or rex CHCEPEPEREELECCT ESTEE 33
ALCOCK OS REAR OAT rele meaileenia ese UE ta. 36
DOUV ICT EON NE OTERO reac scette ee 33
CHE ee rae saute tes 33
COCN LID ES ees ceeastaettntedetieotncsestesstess 33
COOPEN Ie rneiacecetesrerteeietie nek cond tie oat tee socrtoreei nner 33
COT UD A oe een ercagasacewussventeiciscsssaseccesee 5; 34
CAT Desereecs tis pierces a ventrrrry ete choc fla CaP iey ae 34
DART A ESET IN SUNG Ue Nv acticeeess 34
AAS HON ellatiteiibabiatcbeeeluidhuncehneeland te hs 5; 34; 76
DOME PIMA AA eee ee eo sa 34
CAW ANSI OOM CNL hectte trem nee. 34
ID GN INCL Tete eee nL race canteen ota 36
IB QLGVARENISIS) <. scce jecccunisstectattesneenteeassssedesrcaceus 4; 34
(QLOWLCNISUS Soon cscccts tes cctesutiaaiocnstessscsctaavccéecéiesessi ses 34
IST ACIIpe swtorrderesternere aenntemetetinrsen rere acer neste Sh)
DUC QA eect asa var Nestacnrascsieiesursoeaehytoestessecnesros 3; 35
TGACHODUS Hee siete eRe dae ene seoe eons 35; 76
TRACT OSPINU CV Apne 35; 98
ITUUS GUL i axccne tee ear ease ie 35; 77; 78
UL STCNISES ee sois cetera saseoe sesctstesevinvates Suscshecshiis 35; 76
PHIL PPINENSIS scvccveasessccevaivenssevessvexessesiwseseseee 35; 76
OTST CG ore et peer Ea Tec CEE Coe CREE EERE 35
POLL ICUG [EAS Bie beseccostec oc eee eo eae 36
PTINING odes sawoaduatscouscwatscvessseaeess hee cok ee 36
QUGCKULODGIG ei eeirereci treat ee ise ae 5; 36
SULA recator pesca ane edrasaraeal eee Netseseutsees Oe 4; 36
ISPD ERA EV, aa eee actee taeda cesar eee eae tan eices ese Sees 7; 36
WOOAMASONE ircce sec ccccenciicsccocscces oususcse st 36
Thalamitoides Quadridens .0......ccccececctsseetere tener 36
Thalamitopsis, Thalamit ......ccccccccccecssersscssrssesserseeees 34
URGIANONYX\ Or AGIUPES) :secdasescecsvscevsasvn tee 35
ihalammony x WUAGIGMIN Gs. .5.0:scccscecsoesssterewecesevendsresceurs 35
ThalassograpSus Rar pax ...ccccccicccscscesereseceneieess 69; 77
WU RALASSUCA SOCAN een ene aale onc crete Sucdeicereasetcesite 74
Thelphusa
UBC ATID eres eee A EUR CeO DEES DEO 77
WULLLERSEONE Soe cavisicesecscten tt estan eae ees ohee oles Wf
theresae@HOGnee’h ochicciatasissateveiesssisttiivinsessutieiceeen 10
thukuhar, MetopOgrapsus ....ccccccescesersescssteceesesenseeess 68
HDICenNEGICINUS). cert icrecas sc esiisccste cas tseescsteseectevicsess 14
PIQUING MUN GDEZIG sesescscsessestscessoonsivve dt ester sa teventeetes 5; 60
tomentosa
JIN ELTAT ERT] Oye IA Use Es aE EO ICN SR RE a 43
113
I DREGE I GTA ccececoe eacHoPECOLEE racer EEA DOSAESCCOTEPEEEE DED 22
tomentosus
A CIACOd CS trreer rae eecc ran tact naan aetna ee re gel 43
PA GIUITITIUS Mes nea eae casas eaios eects oa ae eens Meee 61
REL OUSINES ere eee oS 22
transversus, PAChYQrApPSUs .........0.cccccceseesecnscsscesseees 76
Trapezia
CN COLAE Tee ee rete eset ee Ted bekeyal See 3; 58; 60
Del gener laa i iene istraK rk coat tees nls BN 3; 58; 60
CYMNOUOCE NE ater ee 58; 59; 60; 61
CLAVAOENSIS ee eR Ean Nore es 60
ONL UN theese Gesu aeseokoeetab aeesneas Meter 58
AIG ital is OOO AR RELL hort te 58; 59; 60
SENT UB INCOR ster cise cere ete I, 58; 59; 60; 61
PLAVOPUNCLAIG Bek neti csecete reece neces cee eoetes 59
fOPMOSO et ss RR She 59;77
USCA EB ei HE ee Get taal 59
DULL ALG as aa caecasvesesscateassextg snes estasitosseduaeseedess 58; 59
WALT OR cece c eee OC ee SEP ENCE EO PCE REE OPP E Ter 58
TQCULGL A re eae ER Nay en Sots UO
LAL ICLIZTIC Is nas boone soa a obo EE eSoC HOOD OLED EAS SOT COoEr 59
PUNCUMONUS esse) Scsecncascusteckscatsenresusesessass 6; 58; 60
FEUCUIAIG cee eS 58
Uf OPUNClalapeennnaes ae ee 59; 60; 61
SCD Ginrairseiesin oa: 58; 60; 77
SCT ENE Liha en arene eRe eh RTA oak Byes 6; 59: 60
SPECIOSA si. oe ern ee 58; 60
HBFING shi eee 5; 60
WON Cth ni mmtoreta beta teal ees leek ua, SESS NENENAN, SONNY 61
UI; ADEZOIUCANSESAVING renee eee eee 70
trapezoideum, LaAbUGnium .........c.cccecceeseetecnteeee escent 70
MEQUIETA WEUMUNIA deserere eects eee eee 95-97
LFiGAGHINALUSNOZIUSE ee 64; 78
Trichopagurus trichophthalmus .......1..100cccccsceeeees 19
trichophthalmus, Catapaguroides, Trichopagurus .. 19
tridens
Caphyrapecncanen ee 30
Gryplodromiopsisnecen: cates ee 24
IFISPINOSUS) SYIMPAQUIUS).caccsccxccoucetevesscneveerre nee seeere 96
tristis
Garpilodesteccvaceenkencct RO ES: 40
NOME Brie raskatc coe covdeveseeecctusoewiooiactenazyueatae 3; 40
Priunguiculatus, JONESIUS .........ccceccencesseeceseeessceeneenseee 3/
UF IZOPACUTUSTSULE ALU Serna nent ere eteeeneee ee eee 16
UNG ALT OMSWIEIDY SLES eereececesc asec seeroereen start cuentees 29
UEUNCALUSROZIUS Ser ho eats orc asre oe eek oes 65; 77
LUGMOLENS OCHRE Girccscdick koscexessdavccc tee 98
LM AMOLUN SYMP AR UTUS | ess ceccecseccse en -sooeettetaeentet se ktsees 96
tuberculata
(CONE 0 cease OER EEOC EE CREE SORE ck 25)
LAR USI isos ieescuuceusdussote.saveicauseidvae Rapsc MN ee 72
tuberculatus, Menaethius ...........c0:0cecceceececeeceeceseeceeees 27
LUDENGUIOSUSROZUUS ernie aoe cheer ee eeee TE 65
EUITIA ARICA EQ spo eee ooo earch e aes 27
tumulosa, Actaea, Paractaea, Paractaeopsis........... 44
Tweedieia
LAY SQL Foci easels cakes sla ceases coveted ganaacedesguscees chess 57
ODDO NU a Oe ares cian veces vateleg ieacs ous vaiaaweasesipisiceseses 7,
Tylocarcinus
UIGHUR Rea c ao eck cecaee nets caste oak capSbnke etek canes 28
OT ACILES cos aatrensn ue acessatevoneasvevesesclas sSscsouecteecsaaneets 28
Ti LOU DION ISP ass occ otoccesccan so xasucaes (Svctgncsibexcéseteustcueers 99
EY DIGUSWANIIGULUS seccecckestacescvans ses sotsdcecesevscsrcseseeceewct sss 13
114
Uca
Ghlorophinalmswrniccsicees eee 74; 80
CUHOUSSI OS boo ours ase Poza ssta sense epasapta stone Sepnaxte 74
GU SSUNMONR Uc Bae saree, co cease orci sesen ctor nite 74; 76
ID CIINGT. GUY, cos cer weatsh ates bee decay sseatstiseeae doctacetese tee eaee 74
SUI OM ee sis cssaacs sass sncsutesssessioessaccpebaate ciusasoea> 74
VAT AITO CRO ee 3; 74
ungulatus
(DREADS eee oe 5)s)
TAN IIOOIUS a rons cassie cones tease issuvenevas coc agstaaNameesentec’ 55
TAI COLIULUS HE GIIPUESLUS) csscessscereensoonseeedeteeeeeeeees eae 96
MEP SUIS= IG] OFS PGI TE DACUS watever ace ner se) cece eet iH
VINCI OCY DOE, ti cscccscccevesassussscxcscreceves uaceceeeeemagi: 73
V
vaillantianus, Carpilodes..........ccccccecensessesensecseeeenee 38
VI ARUMIN GNI OIE) .csic:.0.002<0s0ess <0 .005-52 SAR 98
Variegatus, LEptOgrAPSUS .........cseccsercvecssvscsetessevenees 67
variolosa
INC OLIOINEN, GiB scoters. sas. dose sca nessessanesee ius SEREOS 41; 45
IP SCH OMOMEN.G sox. Hoedscvoegees seakesaccessio1sesstueeeeees 45
WASTED TTR CG Mey perce e oo a Reon Re SSSOACOO ho 6; 69
VEMOSC WA OMENG ics ssicagasxsnaasacisaad ree Re 40
VENOSUS, |G At PULOd OS). .nnn:-----t8so esses ess ec Nes 40
venusta
(QUO Od OP SIS 5 cio sonia ncvensosornnesctaseeae ous ae ees 5/7,
CLE ONU CIO éics50653 SR A eR eS 26
Verdensis;|GarGiiOp aX a vsasaccosesteees to. eect eae 98
VETIGELIPHUTESTELE Qi tenes. <i irck Cina wane scocheatecs Sacto en 96
VERSIGOLOM TE CHUUTUS teen ecet ee ee 10
WIGAEN IN OLOSCELES:. «ns 540<040 0s sss ce RT eee 97
WECLOM VIGIL Gres sscocSccccssteos tour tectete ed he RO AYU
Vi Gil POAOPRURGLINGS ......11...s0+200.nevecn-cusossaesecersan<thees 37
VILLOSUS: (ROLVACCLUS) <a ccscnseschs.canscnsus-ncsss-seneaeee- Sif
Ww
WALLSE SNIP AR UBUS) «icncascsseasussoaciaecaadateres See ED 96
Wardl, L1apezbajsk ss stttess Mapcterc SER, Ne 61
WICHEGKI, EGIPUTELINS cc.sscecsceracsescescsocnsceeeek 10; 76; 77
wilsoni
DONA ied os Latch ee eek 7; 24; 97
PetQlomMen ds on cecisncncsacecscs RO ES 24
WOOGINGSONY: THGIGUINGnceessncss<1scsae eee ee ee 36
WILEDSLONSE, LIMEIDMUSG ooa:-2....2eseeestaets oats eee 77
X
Xanthias
RTL TEL LT EET EGTA TIN pee oy ee en RR ELIS 50
Lig ATL RG) fly rey eR ee te RR 50
VES /V OFS. Pe so cce econ ests sna So Spannacevackiseassoe-oe sea 50
IAEA TAATLETRS coc ty Sa a De 50S GF.
WLOLAUIS vox, Shands soon tec vs asec sooo seated vaeaa has ee 50
JP OMADENSIS ices cies ccussesssteesesusassdsesecncesentouesSeuagteeeeee 48
DUNCL GUUS Vii Soca.ccs0t ccssessvsenstinesseerctesonscseccacanaeeaeer 51
Slee ecto este Ree oPh OSE coco seanee cence tanceanetee 99
LCL BOD ORO ikea cescssulecs bivesavcessencevodecppeeeen SLIT
Xantho
QU OCUGIUS Eo ccceccsnccssvasssvananaspiptoanshee Bosal oe 48
CK GSSIINGIUS) Pi occcacdccasteccccatsstzastheaeeeceeee ee 49
CLAN GUSH Ak svsngattsscsdcsooeeters seks ee 49
OH AGUS Ee 20005, sa. sncdnsiacsccaccenssuesasieseuscssteacceaneeaeae 49
SQN OU ICUS hoi cia scacvas coveseacanssuncesesttssanteeege career 49
SUDET DUS ree casos Sancancvesisndecsueseusa:sseeee eee 46
EGIL CNISI SPREE: «505a jesse deecseasseesoesoenavesneen SRR 48
BCL AOD ONE RBE. casa sccsenstacsacassaanannte ence eee 5]
Xanthodes
RT ANOSO=INQNUS |i s0i0.c<s00sessoseensenxcosesnorenueeaneaeeeeee 50
TANLECL ULL S ORK oo 22 «ca cae nancaccanadeu=cesoee sate ee 50
POLLS AIEEE oa Sans scbev ane susenvew eee 50
AN SUOGUUISICTESIGLUS ¢<s06sass<seasevas saecesnccccsess eee ee 47
xanthoides, ZOZYMOAES ...........+1-c1eereeeerseneee 47; 48; 77
PX PHONECIES. POTLURUS oonceccencscureaccosoevnsseanencnseean 31532
Y
Wald WyNOpSiS SPUN GRUS). «.2...2<<.0c0cpexattendsencssntoenctoaee 97
Z
ZED Gs GUDANATIUS scsccskccoscaseeccancseonseupne senso TOO 16; 17
Zozimus
LO TIOUIN cetacean cad ascgacauad canada SRAM NARS RR 47
SPCR ES ats cxasnsiasenaanerennonnstiedaca aaah Oe 47
Zozymodes
COLINI DOS as saccnsneonsnascnsnnss SSR RD 47
PUIAULUS sorsssoasseccosouneossnnseTedeeeetaeaste to MeReueN ages 47
KANUNOLES, eiosecasansaciceccavcnnns SORES 47; 48; 77
ATOLL RESEARCH BULLETIN
NOS. 435-442
NO. 435.
NO. 436.
NO. 437.
NO. 438.
NO. 439.
NO. 440.
NO. 441.
NO. 442.
MORPHOLOGY AND MARINE HABITATS OF TWO
SOUTHWESTERN CARIBBEAN ATOLLS: ALBUQUERQUE AND
COURTOWN
BY JUAN M. DIAZ, JUAN A. SANCHEZ, SVEN ZEA, AND JAIME
GARZON-FERREIRA
CORAL FAUNA OF TAIPING ISLAND (ITU ABA ISLAND) IN THE
SPRATLYS OF THE SOUTH CHINA SEA
BY CHANG-FENG DAI AND TUNG-YUNG FAN
FIRST OBSERVATIONS ON THE FISH COMMUNITIES OF
FRINGING REEFS IN THE REGION OF MAUMERE (FLORES-
INDONESIA)
BY MICHEL KULBICKI
GROUPER DENSITY AND DIVERSITY AT TWO SITES IN THE
REPUBLIC OF MALDIVES
BY ROBERT D. SLUKA AND NORMAN REICHENBACH
EFFECT OF TYPHOONS ON THE LIZARD COMMUNITY OF A
SHELF ATOLL
BY MICHAEL JAMES MCCOID
FLOWERING AND FRUITING IN THE FLORA OF HERON
ISLAND, GREAT BARRIER REEF, AUSTRALIA
BY R.W. ROGERS
NAMU ATOLL REVISITED: A FOLLOW-UP STUDY OF 25 YEARS
OF RESOURCE USE
BY NANCY J. POLLOCK
CRUSTACEA DECAPODA OF FRENCH POLYNESIA (ASTACIDEA,
PALINURIDEA, ANOMURA, BRACHYURA)
BY JOSE?H POUPIN
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
APRIL 1996
ATOLL RESEARCH BULLETIN NOS. 443-449
>
See re
RESEARCH
BULLETIN |
Issued by
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. U.S.A.
OCTOBER 1997
ATOLL RESEARCH BULLETIN
NOS. 443-449
NO. 443.
NO. 444.
NO. 445.
NO. 446.
NO. 447.
NO. 448.
NO. 449.
THE EVOLUTION OF A HOLOCENE FRINGING REEF AND
ISLAND: REEFAL ENVIRONMENTAL SEQUENCE AND SEA
LEVEL CHANGE IN TONAKI ISLAND, THE CENTRAL
RYUKYUS
BY H. KAN, N. HORI, T. KAWANA, T. KAIGARA, AND K.
ICHIKAWA
CHECKLIST OF THE SHOREFISHES OF OUVEA ATOLL, NEW
CALEDONIA
BY MICHEL KULBICKI AND JEFFREY T. WILLIAMS
ON THE ORIGIN OF DRIFT MATERIALS IN THE MARSHALL
ISLANDS
BY D.H.R. SPENNEMANN
DISTRIBUTION OF RAT SPECIES (RATTUS SPP.) ON THE
ATOLLS OF THE MARSHALL ISLANDS: PAST AND PRESENT
DISPERSAL
BY D.H.R. SPENNEMANN
A POSSIBLE LINK BETWEEN CORAL DISEASES AND A
CORALLIVOROUS SNAIL (DRUPELLA CORNUS) OUTBREAK
IN THE RED SEA
BY ARNFRIED ANTONIUS AND BERNHARD RIEGL
MARINE ALGAE FROM OCEANIC ATOLLS IN THE
SOUTHWESTERN CARIBBEAN (ALBUQUERQUE CAYS,
COURTOWN CAYS, SERRANA BANK, AND RONCADOR BANK)
BY GUILLERMO DIAZ-PULIDO AND GERMAN BULA-MEYER
SCIENTIFIC STUDIES ON DRY TORTUGAS NATIONAL PARK:
AN ANNOTATED BIBLIOGRAPHY
BY T.W. SCHMIDT AND L. PIKULA
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
OCTOBER 1997
ACKNOWLEDGMENT
The Atoll Research Bulletin is issued by the Smithsonian Institution to provide an outlet for
information on the biota of tropical islands and reefs and on the environment that supports the
biota. The Bulletin is supported by the National Museum of Natural History and is produced by the
Smithsonian Press. This issue is partly financed and distributed with funds from Atoll Research
Bulletin readers and authors.
The Bulletin was founded in 1951 and the first 117 numbers were issued by the Pacific Science
Board, National Academy of Sciences, with financial support from the Office of Naval Research. Its
pages were devoted largely to reports resulting from the Pacific Science Board's Coral Atoll Program.
All statements made in papers published in the Atoll Research Bulletin are the sole
responsibility of the authors and do not necessarily represent the views of the Smithsonian nor of
the editors of the Bulletin.
Articles submitted for publication in the Atoll Research Bulletin should be original papers in
a format similar to that found in recent issues of the Bulletin. First drafts of manuscripts should
be typewritten double spaced and can be sent to any of the editors. After the manuscript has been
reviewed and accepted, the author will be provided with a page format with which to prepare a
single-spaced camera-ready copy of the manuscript.
COORDINATING EDITOR
Ian G. Macintyre National Museum of Natural History
MRC-125
ASSISTANTS Smithsonian Institution
Kasandra D. Brockington Washington, D.C. 20560
William T. Boykins, Jr.
Theodore E. Gram
Jonathan G. Wingerath
EDITORIAL BOARD
Stephen D. Cairns (MRC-163) National Museum of Natural History
Brian F. Kensley (MRC-163) (Insert appropriate MRC code)
Mark M. Littler (MRC-166) Smithsonian Institution
Wayne N. Mathis (MRC-169) Washington, D.C. 20560
Victor G. Springer (MRC-159)
Joshua I. Tracey, Jr. (MRC-137)
Warren L. Wagner (MRC-166)
Roger B. Clapp National Museum of Natural History
National Biological Survey, MRC-111
Smithsonian Institution
Washington, D.C. 20560
David R. Stoddart Department of Geography
501 Earth Sciences Building
University of California
Berkeley, CA 94720
Bernard M. Salvat Ecole Pratique des Hautes Etudes
Labo. Biologie Marine et Malacologie
Université de Perpignan
66025 Perpignan Cedex, France
PUBLICATIONS MANAGER
A. Alan Burchell Smithsonian Institution Press
ATOLL RESEARCH BULLETIN
NO. 443
THE EVOLUTION OF A HOLOCENE FRINGING REEF AND ISLAND:
REEFAL ENVIRONMENTAL SEQUENCE AND SEA LEVEL CHANGE IN
TONAKI ISLAND, THE CENTRAL RYUKYUS
BY
H. KAN, N. HORI, T. KAWANA, T. KAIGARA, AND K. ICHIKAWA
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
OCTOBER 1997
35°N
KYUSHU
f
Northern Limit of
Yr _. Coral Reef Formation
+ 30°N
| BASTERN
CHINA SEA Ke #
{\
Tonaki Is. Fig.2A / °
SS
’
25°N
130°E 135°E
Figure 1. The Ryukyu Islands, an island arc at high latitude for reef growth.
THE EVOLUTION OF A HOLOCENE FRINGING REEF AND ISLAND:
REEFAL ENVIRONMENTAL SEQUENCE AND SEA LEVEL CHANGE
IN TONAKI ISLAND, THE CENTRAL RYUKYUS
BY
H. KAN!, N. Hori2, T. KAWANA3, T. KAIGARA4 and K. ICHIKAWA>
ABSTRACT
Within the Indo-Pacific region Holocene reef development over the last 6000 yBP
has occurred during a near stable sea level period. In particular, development of reef flat
and related features have been associated with the stillstand. This is illustrated by the
Holocene evolution of a reef in the Japanese Ryukyu Islands. A continuous reef
structure, 8 m in thickness and 630 m in length, was observed from a fresh excavation in
a modern fringing reef in the western part of Tonaki Island of the central Ryukyu
Islands. Documented by 34 radiocarbon ages, the reef first reached a relative sea level of
ca. 1 m above the present level at about 5200 yBP by growth of branching Acropora
thickets, and by accumulation of angular clasts of tabular Acropora at the landward side.
Storm features occur within all the observed reef structure and suggest that the reef has
been continuously affected by high-energy events for at least 5500 yBP. The
topographic outline of the modern fringing reef was formed at an early stage of sea level
stillstands affected by wind, climate, and substrate topography. Delayed closure of the
‘Holocene high energy window’ resulted in vigorous reef growth on the landward side.
The reef flat accreted seaward about 400 m during the last 4500 yBP with the
development of spurs and grooves. The growth environment shifted from a sheltered to
a wave-affected condition during the seaward accretion of the reef flat. Reduced colony
sizes of tabular Acropora and decreased upward reef growth rates also occurred at about
this time. With the seaward accretion, the shoreward grooves became isolated and
infilled by rounded clasts. Radiocarbon age from an early archeological feature (Touma
and Oshiro 1979) indicates that the formation of a Holocene tombolo is closely tied to
sea level fall around 3500 yBP. This sea level fall and sufficient development of wave
resistant structures at the reef edge, provided shore protection and contributed to
tombolo stability.
! Department of Physical Geography, Faculty of Education, Okayama University,
Okayama 700 Japan. 2 Department of Geography, Faculty of Science, Tokyo
Metropolitan University, Minami-Osawa, Hachioji 192-03 Japan. 3 Laboratory of
Geography, College of Education, University of the Ryukyus, Okinawa 903-01 Japan.
4 Department of Geography, Kansai Gaikokugo University, Makikata, Osaka 573
Japan. 5 Department of Geography, Komazawa University, Setagaya 154 Japan.
Manuscript received 2 September 1997; revised 24 September 1997
INTRODUCTION
As coral reefs grow they can alter their own environment. In mid-Holocene times,
reefs that were still catching up with sea level had minimal wave baffling effects.
Neumann (1972) named this the ‘Holocene high energy window’. Subsequent reef
growth produces a protective crest which in turn results in the development of the back-
reef area. On fringing reefs, in particular, this may result in a shallow lagoon or moat
with ponding of terrestrial run-off which may limit coral growth (Ginsburg and Shinn
1964; Schlager 1981; Neumann and Macintyre 1985; Acevedo et al. 1989).
Sea level stillstands of the order of millennia can produce substantial fringing-reef
flats (Buddemeier and Hopley 1988). In the Indo-Pacific region where a Holocene
stillstand has occurred for approximately the last 6000 years, understanding of reef
growth in response to both early Holocene sea level rise and late Holocene stillstand is
needed to fully understand reef evolution.
This study reports on the evolution of the modern fringing reef and sea level
change in Tonaki Island, the central Ryukyus (Fig. 1), as deduced from a continuous reef
section in a harbor excavation across western Tonaki Reef. Supporting evidence also
comes from coastal landforms including a fossil reef surface overlain and protected by
beach conglomerate, marine notches, and a tombolo (Fig. 2). Tonaki Island is a high
island with two small hills consisting of late Paleozoic and Cenozoic formations
(Konishi 1964) joined by a tombolo (Fig. 2B). The tombolo is 330 to 740 m wide and
330 to 1000 m long. Some of the features of the tombolo developed on Tonaki Island are
similar to the cay islands of the Great Barrier Reef (Gourlay 1988), Solomon Islands and
British Honduras (Stoddart 1969).
Hopley (1968; 1971; 1975) described development of island spits on continental
high islands of North Queensland and concluded that the many similarities between the
fringing reefs and associated deposits of the high islands and the low wooded islands
(complex reef islands) implied a comparable Holocene history (Hopley 1982, p371).
Reef structure and radiocarbon dates observed from the western Tonaki trench and
coastal features has enabled us to discuss the time-series relationship between reef
environmental change, sea level change, and subaerial deposit formation by reference to
Figure 2. (A) Location; (B) general topography of Tonaki Island. The contour lines are
drawn at 50 m intervals for land topography. Numericals in B indicates sampling sites
of dated corals and beach conglomerate shown in Table 2. Location of observed reef
section is indicated in B and C. (C) Modern reef topography of the northwestern
Tonaki Reef. Bold arrow shows submarine ridge extending to the northwestern small
rocky island (Irisuna Island). Reef zonation: a reef slope (J furrowed platform, 2 with
spurs and grooves), 6 spur and groove zone of reef edge, c reef crest, d inner part of
reef crest (some area recognized as a rubble flat), e shallow lagoon (3 coral alignment),
f inner reef flat, g beach (4 beach rock), h land.
WONARS
ISLAND
i
archeological ruin
(Higashl shell heap)
c
RS,
observed tombolo *
reef section (Fig.3) : oy
{3 .¢ Zamami
soNL 1B.
Vie. a te
Me A om Hy
Yi aoa)
| ) ON WGE Oe
3 4 j y Pf 2) 0 As
3B Z NGSG8
Bae,
.’ + g& observed reef section
MW gs 9
4
these observations and radiocarbon dates of a fossil reef surface and the earliest
archeological ruin on the tombolo of Tonaki Island as described by Touma and Oshiro
(1979). This relationship between the reef and associated Holocene landforms indicates
how reefs respond to external and internal environmental change.
MATERIALS AND METHODS
In the Ryukyu Islands, many trenches have been excavated to construct harbors
and associated ship channels across modern fringing reefs. Such excavations provide the
opportunity to observe the continuous structure of modern reefs. Reef structure
directly observed from trench walls reveals the exact location of sedimentary facies,
shape and size of coral colonies and coral clasts in contrast to observations made from
reef cores (Kan and Hori 1991). In Tonaki Island, a harbor excavation reaching 8 m in
thickness and 630 m in length across the western Tonaki Reef, provides a continuous
reef section (Fig. 2). The southern wall of the excavation provided a fresh reef section
because the survey was carried out immediately after widening of the trench in 1989.
The reef profile (Fig. 3) was surveyed using measuring tapes, a 5-meter pole and
depth meter to define micro-topographic features and to provide the basis for later
descriptions of the section. Descriptions of the reef structure were carried out from the
reef surface to the foot of the excavation at approximately 5 to 10 m intervals along the
entire length of the section. The survey was carried out at closely spaced horizontal
intervals especially at the boundaries of the sedimentary facies. Fossil corals were
collected from the excavation by using a hammer and chisel. Twenty-five radiocarbon
ages were obtained for corals in this reef section (Table 1). These dates were established
by methanol liquid scintillation counting, at the Dept. of Geography, Hiroshima
University. Laboratory procedures were based on Fujiwara and Nakata (1984).
The geomorphological map of the northwestern Tonaki Reef (Fig. 2C) was
prepared to document the zonal and the micro-topographic features. The reef
topography is based on the interpretation of field observations in 1989 and the 1977
color aerial photographs scaled to 1:10,000. Coastal landforms such as fossil reef surface
overlain by beach conglomerate, and marine notches were investigated to document mid-
to late-Holocene sea level indicators in 1991 and 1996 (Fig. 2). The altitude data were
revised with reference to tide tables. Tidal values in Tonaki Island are corrected by
reference to those in Zamami Island, near Tonaki (Fig. 2). Mean high (low) water level
in Tonaki is about 0.6 m above (below) mean sea level (MSL) which is 1.16 m above the
tidal datum. Tidal range is approximately 2.0 m at spring tide. Nine radiocarbon ages
were obtained from in situ corals in the modern reef surface and in the fossil reef surface
overlain by the beach conglomerate (Table 2). These ages were dated by Prof. emeritus
K. Kigoshi of Gakushuin University.
X-ray diffraction was used to confirm the absence of calcite in all dated samples.
Age calculations are based on the Libby half-life of 5568 + 30 years. Errors are indicated
at the range of + 1 o. The dates have not been corrected for isotopic fluctuations or
5
environmental factors because we have no data on the ocean !4C reservoir effect for the
Ryukyu Islands.
RESULTS
Topography of western Tonaki Reef
On Tonaki Reef, a reef crest has developed in the north where topographic
zonation is clearly defined, and flat topography with poor zonation occurs on the
western reef (Fig. 2C). On the northern part of the reef, spurs and grooves on the reef
edge and furrows on the submerged platform off the reef edge, at a depth between 10 to
15 m align with the prevailing northerly wind of winter seasons. These features are
similar to the northern reef of Kume Island described by Takahashi and Koba (1977)
and Kan (1990), which lies 30 km west of Tonaki Island. The distribution pattern of
reef slope coral ridges changes abruptly on the northwestern reef slope where a
submarine ridge extends westward (bold arrow in Fig. 2C). The long axes of the coral
ridges and honeycombed furrows suggest that a southwesterly current dominates on the
western reef slope. The observed reef section bisects a large number of spurs and
grooves (Fig. 3), because the orientation of spurs and grooves is oblique to the reef edge
on the western reef (Fig. 2C).
The dominant coral assemblage of the reef edge is tabular or encrusting Acropora.
On the northern reef, a coral rubble flat of tabular Acropora clasts has developed behind
the reef crest. In the shallow lagoon, branching Porites dominate in the northern part.
However, the lagoon tends to shallow towards the south where sea grass beds, with
branching Montipora and Pavona cactus, are found.
Internal structure beneath the reef flat
Four major bio-lithofacies are identified for the upper 8 m of reef structure in the
western Tonaki reef flat. Three distinct zonal structures were observed: from seaward,
tabular Acropora framework facies; tabular Acropora transported rubble facies; tabular
Acropora reworked facies; and branching Acropora facies. These facies are arranged
vertically in the reef structure (Fig. 3). The other lithofacies described below are
subordinate.
1) Tabular Acropora framework facies: Jn situ growth and accretion of tabular or
plate Acropora dominates the outer half of the reef structure. This facies constitutes a
large part of the convex topography of spurs or paleo-spurs (Fig. 3) and abut thick beds
of tabular Acropora transported rubble facies described below (Fig. 4B). While large size
in situ tabular Acropora colonies, around 2 m in horizontal dimension, are dominant on
the inward end of this facies zone (Fig. 4A: 340 m point in Fig. 3), colony size decreases
to 20 to 50 cm at the outer edge. Coincidentally, the upward reef growth rate gradually
decreases seaward (Table 1) as follows: 8.2 m/ky (between TN-12 and 13) for the earlier
spur dated around 4500 yBP; 4.1 m/ky around 4000 yBP (between TN-6 and 7); 3.6
6
m/ky (between TN-4 and 5); 3.2 m/ky (between TN-2 and 3); 1.2 m/ky (between TN-1
and present).
2) Tabular Acropora reworked facies: The unconsolidated platy clasts of tabular
Acropora have accumulated (Fig. 4C) to at least 6m thickness and formed a sedimentary
zone behind the tabular Acropora framework facies (Fig. 3). These Acropora clasts (Fig.
4D) are relatively well preserved and essentially overturned in situ colonies. The
radiocarbon ages between TN-20 and TN-21 are reversed, despite 2.6 m difference in
their depths (Table 1).
3) Tabular Acropora transported rubble facies : In the outer half of the reef
structure, rounded coral clasts (Fig. 4E) have densely infilled the in situ tabular
Acropora facies (Fig. 3). Some clasts are coated by calcareous algae (in the form of
rhodoliths; Fig. 4F), which suggests that they had been tumbled. These are similar to
those that have accumulated in the present-day grooves of the reef edge. Radiocarbon
ages of rubble lag approximately 400 to 900 years behind neighboring in situ tabular
Acropora facies (between TN-9, 12 and 10; Fig. 3).
4) Branching Acropora framework facies: Thickets of in situ branching Acropora
(Fig. 4G) form a framework zone which exceeds 100 m in width and is more than 6 m
thick (Fig. 3). These colonies were relatively intact (Fig. 4H). However, calcareous algae
which covered the surface of branches makes species identification difficult.
Radiocarbon ages (Table 1) show this coral thicket had been growing for over 400 years.
The observed upward reef growth rates were 6.3 m/ky (between TN-14 and 15) for this
thicket. Aside from the fine sediment that covers the excavation (Fig. 4G), little
sediment was observed infilling the branching framework.
(TN-2) (TN-4) (TN-6) (TN-9) (TN-10) — (TIN-12)
880-55 2220+60 3900+65 3890 +70 3510+65 4390+70
—=— Seaward
Om MSL
---- tidal datum ----------- 3
(TN-1)
-5 2340.60. |
55 pS IL
a? ® ||
eth? yi :
sai Stes 3500-60 11) 4960-65
202060 3390+65 a7) ON) (TN-13)
-10 (TN-3) (TN-5)
100 200 300
= tabular Acropora 2 ©] tabular Acropora
19 framework facies transported rubble facies
tabular Acropora tabular Acropora
0 [= | reworked facies reworked framework facies
Figure 3. Reef structure and radiocarbon ages of fossil corals in the western Tonaki
Reef. Lettered squares indicate positions of photographic sites presented in Fig. 4.
7
5) Tabular Acropora reworked framework facies: Tabular corals and calcareous
algae constitute blocks | to 5 m in diameter. This bio-lithofacies is similar to the tabular
Acropora framework facies of the reef margin. However, directions of individual coral
growth and of accumulated coral colonies are different from the framework facies.
6) Mixed coral framework facies: This facies constitutes corals, calcareous algae
and skeletal grains. No dominant coral genus was observed in this facies.
Beside these bio-lithofacies, the following in situ coral colonies larger than 2 m in
diameter are identified in the growth fabric: foliaceous Heliopora, stubby branching
Acropora, and hemispherical Lobophyllia. No terrigenous facies or Pleistocene limestone
was observed within the section.
Late-Holocene sea level change
On Tonaki Island, part of the Holocene reef flat is surmounted by undercut
blocks (Konishi 1964). It has been suggested that the reef flat originally developed 80
cm higher than the present sometime in the late Holocene (Konishi e¢ al. 1974). Beach
conglomerate which overlies the landward end of the reef has also resulted in protection
of the reef surface against erosion (Fig. 5). These features, together with marine notches
cut into the Paleozoic limestone cliffs are considered to be sea level indicators (Kawana
1996). Nine radiocarbon ages for corals obtained from the modern and fossil reef surface
(Table 2, Fig. 5) show that the sea level achieved a maximum ca. 0.9 to 1.3 m higher than
present ca. 5200 yBP. This mid-Holocene maximum level extended until around 3650
yBP, with a relatively sharp fall taking place at that time (Fig. 6).
(TN-14) (TN-16) (TN-17) (TN-20) (TN-22) (TN-25)
4560+70 4600+65 4580+70 5690+75 5190+75 5180+70
-70
Om
} ay my wa Ae GS Se
x a , mn BA ool Aue we Ne v2 ¥
Pf er ee ee eee e 5
aN = Seek OU ae
Qe A a ONT an ae
ON Ae ones © ye
za 5510+75 3 75
5290+70 Se EMOE) (TN-21) (TN-23) 5660 1
(TN-18) (TN-18) (TN-19) a
-10
400 500 600m
ao mixed coral in situ stubby r~\ invisible portion
framework facies branching Acropora a (wall covered by talus)
branching Acropora | He | in situ foliaceous
framework facies Heliopora
in situ hemispherical
Lobophyllia
bottom of the
excavated wall
dated coral
Table 1 Radiocarbon ages from Tonaki Reef excavation.
Sample Site ae Material Labo. ee ee
oy ae m) ak (yB.P.+1o) Growth
: 1 . :
ao Genus Life Form T( 1/2)=5568yrs het
TN-1 18 6.5 Platygyra hemispherical t HR-561 2340 + 60 - 55 AZ HF
TN-2 70 2.7 Acropora tabular HR-562 880 + 55 32
TN-3 70 6.4 Acropora tabular t HR-563 2020 + 60
TN-4 153 2.1. Acropora _ tabular ft HR-564 2220 + 60 3.6
TN-5 153 6.3. Acropora tabular + HR-565 3390 + 65
TN-6 214 3.4 Acropora tabular ft HR-539 3900 + 65 4.1
TN-7 214 6.1 Acropora tabular HR-540 4560 + 65
TN-8 220 4.6 Acropora _ tabular + HR-538 3500 + 65 - 60
TN-9 280 2.1. Acropora tabular + HR-560 3890 + 70 - 65
TN-10 295 2.1. Acropora tabular %* HR-534 3510 + 65
TN-11 295 4.7 Acropora tabular * HR-535 4210+ 65
TN-12 305 1.8 Acropora _ tabular + HR-536 4390 + 70 8.2
TN-13 305 6.5 Acropora _ tabular ft HR-537 4960 + 70 - 65
TN-14 3)5)// 2.4 Acropora _ tabular t HR-531 4560 + 70 6.3
TN-15 3S 7.0 Acropora tabular + HR-532 5290-70
TN-16 365 3.7. Acropora tabular HR-533 4600 + 65
TN-17 395 2.2 Heliopora foliaceous 7 HR-559 4580 + 70
TN-18 405 5.4. Acropora tabular HR-557 5440 + 75
TN-19 434 5.5 Acropora tabular HR-558 4970 + 70
TN-20 529 3.1 Acropora tabular HR-551 5690 + 75
TN-21 529 5.7. Acropora tabular HR-552 ayo ae ve)
TN-22 555 2.1 Acropora ramose ft HR-553 5190 + 75 - 70 8.3
TN-23 395 5.4 Acropora ramose HR-554 5590 + 75
TN-24 606 6.3. Acropora ramose f HT-555 5660 + 75 - 70
TN-25 625 1.6 Acropora ramose ft HR-556 5180 + 70
* Site shows the sampling location corresponding to the horizontal scale of the reef section (Fig. 2).
+ in situ coral
%¢ rounded clast accumulated in groove
+t Growth rate is calculated on the basis of the TN-1 sample and the top of the spur (present).
The present is assumed, as the spur has not reached sea-level.
Figure 4. Holocene reef structures and components observed from the excavated wall
of the western Tonaki Reef. Sites of the photographs are shown in Fig. 3. (A) Jn situ
tabular Acropora facies, (B) paleo-groove (b) infilled by rounded coral clasts. Paleo-
spur (a) composed of in situ tabular Acropora. (C) angular clasts of tabular Acropora
reworked facies. (D) close-up of angular clast of tabular Acropora. (E) rounded coral
clasts of tabular Acropora transported rubble facies. (F) a section of a rounded coral
clast showing encrustations by calcareous algae. (G) branching Acropora framework
facies. (H) colony of in situ branching Acropora.
10
Table 2 Radiocarbon ages of in situ corals in coastal deposits in Tonaki Island
(see also Fig. 5).
Loc.
No. *
OM YADA HW KR Wb &
Radiocarbon Age
(yB.P. + 1o)
T(1/2)=5568yrs
5150 + 100
4410+ 80
4790 + 140
4430+ 110
4980+ 90
3640 + 190
4130+ 80
4820 + 130
4890 + 100
Material
(Coral Genus)
Porites
Goniastrea
Goniastrea
Porites
Goniastrea
Porites
Porites
Goniastrea
Porites
Labo. No.
GakK-15825
GaK-15826
GaK-15827
Gak-15828
GaK-15829
GaKkK-15830
Gak-15831
GaK-15832
GaK-15835
Explanation for Sampling Site
Elevation of
specimen (m)
above MSL
reef flat - 0.6
reef overlain by beach conglomerate 0.1
reef in front of beach conglomerate 0.1
reef overlain by beach conglomerate 0.2
do. 0.1
do. 0.77
do. 0.4
inner edge of the reef flat - 0.4
reef in front of beach conglomerate 0.2
* Location numbers are in Fig. 2B.
+ The higher elevations of the specimens at Loc. 6 and Loc.7 are probably due to strong wave-
affected coast.
ca. 0.9~1.3m above mean sea level (marine notch)
Beach conglomerate
MHWL
4790 + 140
4890 + 100
MSL
4820 + 130 5150 + 100
3640 + 190
4130 + 80
4410 + 80
4430 + 110
4890 + 90
Holocene coral reef
Figure 5. Schematic profile of Holocene reef, beach conglomerate and marine notch at
the landward end of Tonaki Reef. Radiocarbon ages of corals indicate years BP (see also
Table 2). MHWL: mean high water level, MLWL: mean low water level.
1]
Reef growth and island formation
The western Tonaki Reef first reached the sea level at 5200 yBP by growth of
branching Acropora on the landward side (Fig. 3). The accumulation of the reworked
facies of angular clasts of tabular Acropora was around 5200 yBP. Around 4500 yBP,
some spurs consisting of tabular Acropora framework reached sea level at the outer edge
of the facies formed of tabular Acropora reworked (Fig. 3, TN-14). The transported
tabular Acropora rubble facies gradually became thicker toward the inner reef flat (Fig.
3) where paleo-grooves have been entirely filled by rounded clasts (Fig. 4B). The reef
flat of the western Tonaki Reef has accreted seaward about 400 m during the last 4500
yBP by the development of spurs and grooves. With the accretion of the reef margin,
the landward grooves, which have became isolated from the sea, have been filled by
rounded clasts.
The geomorphological evidence is supported by the history of human settlement
in this area. Touma and Oshiro (1979) excavated the earliest archeological ruin located
on the narrow part of the tombolo of Tonaki Island (Fig. 2B). Trench excavations show
that bioclastic sand accumulated up to an elevation of 3.5 to 3.8 m. Above this sand,
Initial formation
of spur and
groove system om
Development
Reef catch up of tombolo
;
with sea level
—+— ——_— jn situ tabular Acropora ——————_____LLLL_>
SSS eee
tabular tabular Acropora -10
— Acropora transported
-e in situ coral reworked rubble facies
~ tabular Acropora reworked clast facies
-~- tabular Acropora transported rubble oa eae TIS ERT ESS ae Sir a IEC aie CR
Initial formation Secondary
of reef flat progradation
of reef flat
Figure 6. Sea level and reef growth in Tonaki Reef. The sea level curve (solid curve)
until 5200 yBP is based on Kume Island, the neighboring island of Tonaki, when the 3
meters seismic uplift at around 2000 yBP (Koba et al. 1982) is subtracted from the
curve of Kan ef al. (1991). The curve after 5200 yBP is based on the coastal landforms
and radiocarbon ages in Tonaki Island (see text). The dashed lines show the vertical reef
growth at each site in Tonaki Reef (see also Fig. 3 and Table 1). A horizontal bar
indicates the range of error in radiocarbon ages. A vertical bar indicates depth between
the lowest low water level and the mean sea level for Tonaki Reef (0.9 m) to represent
past sea level. The vertical bar is only for ages which are reliable sea level indicators.
WZ
two layers of brown humic sand were observed with a thickness of between 0.3 to 0.7
m below the present-day surface soil. The lower bed, just above the bioclastic sand,
contains earthenware and shell middens. The radiocarbon age of shell material was
3510+90 yBP (laboratory code: N-3080). This age coincides with the chronological
determination of excavated earthenware (Touma and Oshiro 1979).
DISCUSSION
Storm features in the reef development
The Ryukyu Islands are located in the hurricane belt. Catastrophic storm events
result in obvious degradation of coral colonies (see review of Rogers 1993) and produce
large amount of coral clasts (e.g. MacNeil 1954; Hernandez-Avila et al. 1977; Scoffin
1993) during cyclic growth and destruction processes (Done 1992). The fragmentation
of coral colonies appears to be a widespread and extremely important method of
reproduction and distribution (Highsmith 1982). However, overturning of the colony is
critical to survival (Chamberlain and Graus 1975). The morphology of tabular Acropora
colonies which widen toward the top, when broken, produces the overturned
accumulations of the tabular Acropora reworked facies. Storms on modern reefs can
result in deposition in the back reef environment (c.f., MacNeil 1954; Newell and Bloom
1970) and mixing debris of varying ages (Johnson and Risk 1987). The observed age
reversal between TN-20 and TN-21 in the tabular Acropora reworked facies is of similar
magnitude to those obtained by Marshall and Davies (1982) from a drill core which
penetrated into reef flat rubble facies in One Tree Reef on the Great Barrier Reef.
Catastrophic redistribution and abrasion by sediment have been observed during
storm conditions (Kobluk and Lysenko 1992) especially along reef-edge grooves (Kan
1995). The unstabilized rubble is subjected to tumbling (Blanchon and Jones 1995) and
infills cavities (Newell 1956; Edmunds and Witman 1991). The transported tabular
Acropora rubble facies accumulated between the tabular Acropora framework facies
(Fig. 4B) where ‘room and pillar structures’ (Tracey et al. 1948) had been formed. Storr
(1964) showed that water flow over the tops of reefs is erratic and the rate of flow is
generally low, whereas passageways (e.g., grooves, reef tunnels) permit high flow rates.
The initial accumulation site of fragmentated colonies (i.e., back reef or reef edge
grooves) may cause the differentiation between angular or rounded rubble. The observed
reef structure shows that the accumulated coral clasts form rounded rubble (transported
rubble facies) after the development of a spur and groove system (Fig. 6).
Storm events can also destroy reef edge spurs (Stoddart 1962) and throw them
onto the reef flat (Ladd 1961; Newell and Bloom 1970; Bourrouilth-Le Jan and
Talandier 1985). The buried blocks of tabular Acropora reworked framework facies,
along with the accumulation of a large amount of Acropora clasts and the tightly
interlaced branching colonies (as in Fig. 4H) which may also be formed during the
regenerative growth process of fragments (Kawaguti 1937; Gilmore and Hall 1976),
suggest that the reef has been continuously affected by high-energy events.
13
Windward-leeward contrast in the reef development
In the fringing reefs of the Ryukyu Islands, several studies have demonstrated that
the original growth axis, where the reef first reached sea level, forms the present reef
crest (Takahashi ef al. 1988; Kan et al. 1991; Kan and Hori 1993; Yonekura et al. 1994).
Results of drilling in a reef of northwestern Kume Island, located 30 km west of Tonaki
Island, shows the northern reef reached sea level earlier because of shallow substrate and
exposure to the prevailing northerly wind of winter seasons (Kan ef al. 1991). A similar
pattern of formative time lag was described by Hopley and Barnes (1985) for a fringing
reef in the Great Barrier Reef. This pattern of growth helps to understand the
planimetric development of the northwestern Tonaki Reef which has a similar
geographical setting.
A submarine ridge extending on the northwestern reef slope of the Tonaki Reef
(bold arrow in Fig. 2C) may have contributed to the development of the reef crest of the
northern reef because reef development is closely tied to antecedent slope break
(Hubbard 1988; Kan ef al. 1995). The northern Tonaki Reef may well have initially
reached sea level and acted as a breakwater against the prevailing winter northerly winds
(Fig. 7A).
This allowed zonal growth of branching Acropora thickets which is usually found
in sheltered environments (Geister 1977; Pichon 1978; Done 1983). However, branching
coral assemblages in shallow lagoons change in response to burying by sandy sediment
(Nakai 1982). Siltation and suspended sediments also reduce coral growth (e.g., Aller
and Dodge 1974; Dodge et al. 1974; Loya 1976). Mayer (1918) examined 4. hebes
which is a synonym of A. aspera (Veron and Wallace 1984) and same or similar species
to the branching Acropora of the western Tonaki section. It was shown to be sensitive
to the smothering effects of silt. The vigorous growth of branching Acropora for over
400 years, with indications of little sediment influence, may be explained by the tidal
current which may have been flowing between the two islands at about 5500 to 5200
yBP (arrows in Fig. 7A) before the tombolo joined them together.
The northern reef may also have provided the large amount of angular clasts of
tabular Acropora to the reworked facies that accumulated around 5200 yBP, because the
tabular Acropora assemblage had not formed in the western reef but is distributed
abundantly down to a depth of 5 m at the reef edge and decreases in deeper water in the
Ryukyu Islands (Takahashi et al. 1985). At the early stage of reef flat formation,
development of leeward reefs is strongly affected by the windward reef formation.
Styles of reef accretion and formation of reef zonation
Framework accretion with higher accumulation rates is dominated by branching
corals with a high proportion of voids (e.g., Davies and Hopley 1983; Davies et al.
1985; Hopley and Kinsey 1988) and is especially associated with monospecific coral
thickets (Highsmith 1982). Detrital sedimentation by storm events also results in high
reef accretion rates (Davies and Hopley 1983). In the western Tonaki Reef, the early
14
stages of reef were developed by these two bio-lithofacies within a short period around
5200 yBP. They may have also contributed to the sediments of the broad inner reef flat
at the western part of Tonaki Reef quickly infilling any shallow lagoons.
The duration of the ‘Holocene high energy window’ has some regional variation
(Hopley 1984). The development of spurs consisting of tabular Acropora framework
around 4500 yBP (Fig. 3, TN-14) intimates that the ‘high energy window’ has been
progressively closed since 4500 yBP. Subsequently, the area available for active
carbonate production has been reduced to a zone at the reef edge (Stoddart et al. 1978)
as reported in other present-day reefs (e.g., Gladfelter et al. 1978; Kinsey 1981).
Since the size of wave-swept organisms depends on wave exposure (Denny ef al.
1985), the smaller colony sizes of in situ tabular Acropora on the outer reef flat suggest
that the growth setting has changed to a wave-affected condition with the outward
migration of the reef flat. The decline in vertical growth rates at the outward edge of the
reef flat (Table 1, Fig. 6) has accompanied this change.
Contrary to reefs where the crest kept pace with sea level, the delayed closure of
the ‘Holocene high energy window’ resulted in vigorous reef growth toward the island.
Subsequently, however, the area of active reef growth has shifted progressively seaward
with the development of reef edge spurs. This gradual process has contributed to the
build up of flat topography with poor zonation.
Timing of island formation and sea level fall
The time-series relationship between reef island formation and reef development
has been previously described (e.g., Stoddart 1969; Stoddart et al. 1978; Hopley 1982;
Woodroffe 1992). The Great Barrier Reef cays formed after 6000 yBP, coincident with
reef flat formation, but were essentially complete in shape and size by 3000 yBP
(Stoddart et al 1978). On Tonaki Island, the vigorous reef growth around 5000 yBP
appears to have provided the shallow substrate to establish the tombolo between the
two islands and provided enough land for human inhabitation by about 3500 yBP
(Touma and Oshiro 1979; Fig. 7B).
The sea level fall sometime after 3650 yBP in Tonaki Island is quite similar to
those in Kosrae Island in the eastern Carolines of Micronesia where a | m fall has taken
place after 3700 yBP (Kawana et al. 1995), and in the Great Barrier Reef where the mid-
Holocene maximum level may have extended until 3700 yBP (Beaman ef al. 1994;
Larcombe ef al. 1995). Evidence for sea level fall around 3700 yBP appears prevalent in
the western Pacific.
Stoddart and Steers (1977) suggested that reef islands were formed as a result of
sea level fall (see also, Pirazzoli and Montaggioni 1986; Roy and Connell 1991).
Schofield (1977a, 1977b) demonstrated how reef islands formed due to sea level fall and
the supply of sediment from reefs. Sea level fall resulted in reduced wave force at the
shore. The seaward accretion of the reef also contributed to wave attenuation because
15
Pie eet fst Seeeminestt w+ Gar 5000
F igure 7. Schematic diagrams of the formation of the Tonaki Reef. Arrows in Fig. 7A
indicate the “Holocene high energy window’ at around 5200 yBP.
16
waves break farther from the shore. At 3500 yBP, the western Tonaki reef flat accreted
to approximately half way across the present-day reef flat (Fig. 7B). The sea level fall
and reef maturation with development of wave resistant structures at the reef edge,
provided shore protection and contributed to tombolo stability.
CONCLUSION
Sea level achieved a maximum ca. 0.9 to 1.3 m higher than present ca. 5200 yBP in
Tonaki Island, the central Ryukyus. This Holocene maximum level extended until
around 3650 yBP, with a relatively sharp fall taking place at that time.
Early development of the reef in Holocene times was influenced by substrate
topography that produced an initial zonation contrasting the outer windward margin,
and the inner sheltered reef. Spur and groove systems were initiated with major reef
accretion occurring by growth of the reef edge spurs. As inner grooves have become
isolated from the sea, they have been infilled during storms by rounded coral clasts. The
deposition of subaerial deposits of the tombolo of Tonaki Island is linked both to the
accretion of the reef flat seaward, and to a fall in sea level about 3650 yBP.
ACKNOWLEDGEMENTS
The authors are indebted to: Dr. D. Hopley, Prof. L.F. Montaggioni, Dr. M.K.
Gagan and Dr. J. Kleypas for their invaluable comments on this manuscript; Dr. I.G.
Macintyre and Dr. J.I. Tracey for their helpful review; Prof. K. Fujiwara and Prof. T.
Nakata for arrangement to use radiocarbon dating system at Hiroshima University; Prof.
emeritus K. Kigoshi for radiocarbon dating at Gakushuin University; Mr. Y. Nakashima,
Mr. K. Takemasa, Ms. Y. Oka and Mr. T. Toubara for their field assistance; Ms. N.
Nakamura and Mr. K. Yoshihama provide us information about the harbor construction.
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ATOLL RESEARCH BULLETIN
NO. 444
CHECKLIST OF THE SHOREFISHES OF OUVEA ATOLL,
NEW CALEDONIA
BY
MICHEL KULBICKI AND JEFFREY T. WILLIAMS
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
OCTOBER 1997
ae: de ‘Astrolabe
? O
yz
Récif de la Gazelle
: Beautemps-Beaupré G, x
@plle des Pins lle
Walpole
Figure 1. Map of New Caledonia and the Loyalty Islands showing the location of Ouvéa.
CHECKLIST OF THE SHOREFISHES OF OUVEA ATOLL,
NEW CALEDONIA
BY
MUCHEL KULBICKI
AND
JEFFREY T. WILLIAMS |
ABSTRACT
The shorefishes of Ouvéa, an isolated atoll in the Loyalty Islands group of New
Caledonia, had not been surveyed prior to 1990. An extensive survey was conducted by
ORSTOM between 1991 and 1992 to obtain baseline information on the shorefishes. A
total of 653 taxa among 72 families are now documented from this area. The most
diverse families are the Labridae (69 species), Pomacentridae (58 species), Gobiidae (54
species), Serranidae (39 species), Chaetodontidae (31 species) and Apogonidae (28
species). The absence or very low diversity of some families (Clupeidae, Nemipteridae,
Siganidae) or genera (Abudefduf, Neopomacentrus) is similar to findings for other
isolated islands of the Coral Sea. Of the 653 species recorded from Ouvéa, 51 species
have not been reported from New Caledonia, a large high island to the South. Only one
endemic species, Luzonichthys williamsi, has been recognized among the shorefishes at
Ouvéa. A number of Pacific Plate endemic species were recorded at Ouvéa, which is
positioned on the Australasian Plate to the south of the edge of the Pacific Plate.
Antennarius duescus, previously known from three specimens taken at the Hawaiian
Islands, is recorded from a single specimen taken at Ouvéa. Another antitropical
distribution pattern is exhibited by Dinematichthys riukiuensis, which is known to occur
at Fiji, Ouvéa and Queensland in the South and from Okinawa.
INTRODUCTION
Our knowledge of shorefishes in the Southwest Pacific has increased significantly
in the last two decades with the publication of a number of checklists. Shorefishes of the
Great Barrier Reef have been reported from several regions: Russell (1983) and Lowe and
Russell (1990) for the southern part, Paxton et al. (1978) for Lizard Island, Allen (1989)
* ORSTOM, B.P. A5, Nouméa, New Caledonia
** National Museum of Natural History, Smithsonian Institution, Washington D.C. 20560 USA
Manuscript received 4 January 1997; revised 7 July 1997
D
for the Coral Sea, and Paxton et al. (1989: 1 volume, out of 3 announced, of an
encyclopedia of the fishes of Australia, which covers shorefishes). The fish fauna of
Lord Howe, Norfolk and Kermadec islands, the southern limit of shorefishes in the
Southwest Pacific, was surveyed by Allen et al. (1976) and more recently by Francis
(1993) and Francis and Randall (1993). Middleton Reef, a southern reef midway
between Australia and New Caledonia, has been investigated by Hutchings (1988), and
Kailola (1987 a,b; 1991) published a checklist of fishes from Papua New Guinea.
Kailola’s list is being updated by Allen. For New Caledonia there are two checklists, one
for the main island (Rivaton et al., 1989) and one for the Chesterfield archipelago
(Kulbicki et al., 1994; LeBorgne et al., 1994), a group of islands midway between the
Great Barrier Reef and New Caledonia. A number of books are now available on the
shorefishes of the Southwest Pacific region (Fourmanoir and Laboute, 1976, for New
Caledonia and Vanuatu; Randall et al., 1990, for the Coral Sea; Allen and Swainston,
1992, for Papua New Guinea; Allen and Swainston, 1988, for Western Australia).
Despite these recent efforts, the fish fauna of many areas of the Southwest Pacific
remains poorly known. One such area is the Loyalty Islands, which comprise five
islands, Maré, Tiga, Lifou, Ouvéa and Beautemps-Beaupré. The first three are high
islands with very sparse coral reef development. The latter two are atolls, Ouvéa (900
km?) being much larger than Beautemps-Beaupré (120 km?). These atolls are located on
the edge of the Australasian plate near its boundary with the Pacific plate between New
Caledonia and Vanuatu (figure 1), and are the only true atolls within a 1500 km radius.
Only two scientific cruises conducted studies at Ouvéa prior to 1991. Allen, in
June 1973, studied the Pomacentridae of Ouvéa. ORSTOM, a French scientific
organization, organized a cruise to Ouvéa in November 1979, but issued no cruise report
because a cyclone considerably limited their study. A few specimens were collected and
sent to the Paris Museum and the Bishop Museum (Hawaii). We are unaware of any
other samples taken from Ouvéa. Carpenter and Allen (1989) report that Lethrinus
“sp.2” occurs at the Loyalty Islands, but they do not provide collection data for the
material they examined.
In 1991, ORSTOM was asked to evaluate the fish resources of Ouvéa Atoll.
During this survey, data were also collected on the atoll’s physical characteristics (water
masses, geomorphology, sedimentology), the plankton, and the benthos (Chevillon, 1994;
Clavier et al., 1992; Clavier and Garrigue, 1993; Kulbicki et al., 1993a, 1993b, 1994;
LeBouteiller et al., 1993). A summary of these works is also available (Kulbicki, 1995).
The checklist of the shorefishes of Ouvéa Atoll presented here is compiled from data
gathered during the 1973 and 1979 visits and the authors’ sampling program conducted
during their 1991 survey.
MATERIAL AND METHODS
Fish were visually censused along transects and/or collected using rotenone and
SCUBA. Some specimens were caught with handlines. Visual censuses took place on the
lagoon floor down to 25m, and on the outer reefs surrounding the atoll. No censuses or
5)
»)
collections were performed on the outer slope of the barrier reef or on the eastern side of
the main island in the atoll. The locations of the stations are plotted on the map in figure
2. Thirteen collecting stations utilized rotenone (figure 2), the amount of rotenone used at
each station was between 2 and 6 liters of solution containing 8% active rotenone.
20°20’S iL
OCEAN PACIFIQUE
20°25’S L
20°30°S L
20°35’S i
20°40°S L
20°45’S
T ——
168°10’E 1664S’E 166°20’E 166°25E 166°30'E 166°35E 166°40°E
Figure 2. Location of sampling sites (open circles = handline fishing; dark circles =
dives; stars = rotenone).
RESULTS AND DISCUSSION
A checklist, totaling 653 taxa distributed among 72 families, is presented in Table
1. There are about 300 fewer species known from Ouvéa than from nearby New
Caledonia, where more than 950 reef-fish species have been recorded (Rivaton et al.,
1989). The number of taxa known from Ouvéa is closer to the 795 species recorded from
the Chesterfield archipelago (Kulbicki et al., 1994), another isolated area at a similar
latitude, than to the 425 species recorded from Rotuma (Zug et al., 1989), a small cluster
of islands about 450 km north of the Fiji Islands. Because our sampling (only 13
rotenone stations) at Ouvéa is limited and we have not sampled the eastern side of the
atoll, we believe the total number of shorefish species may actually be as high as 800-900
species.
Most of the species in this list have been reported from nearby New Caledonia,
however 51 taxa have not been recorded from New Caledonia. Serranocirrhitus latus
has not been listed in the New Caledonian checklists, but was reported from New
Caledonian reefs and the Loyalty Islands as “ Dactylanthias mcmichaeli” by Fourmanoir
and Laboute (1976). Most of the new records are Anguilliformes, Scorpaenidae or
Gobiidae, which are usually taken only in rotenone collections. There are specimens in
the Ouvéa collections representing 33 taxa that could not be identified to species. Many
of these are juveniles, but 15 of these taxa are currently known to be undescribed species.
Most of the 15 undescribed species are known from other localities. Their descriptions,
thus, will not greatly increase the number of endemic species known from the Loyalty
Islands. The only endemic species described from Ouvéa 1s Luzonichthys williamsi
(Serranidae, Anthiinae).
One specimen of Antennarius duescus was taken at Ouvéa. This distinctive
species 1s unique in having the opercular opening situated halfway between the base of
the pectoral lobe and the origin of the anal fin. It was previously known from only three
specimens from the Hawaiian Islands (Pietsch and Grobecker, 1987). Its presence in the
Loyalty Islands is possibly indicative of an antitropical distribution pattern. Trimma
unisquamis and Gymnothorax eurostus have a similar antitropical distribution pattern,
with a northern population at Hawaii. Dinematichthys riukiuensis exhibits an antitropical
distribution pattern, but has its northern population at Okinawa and its southern
population at the Great Barrier Reef, Ouvéa, and Fiji.
A number of Pacific plate endemic species (Springer, 1982) are recorded herein
from Ouvéa. These records could be described as plate margin occurrences. Pacific plate
endemics taken at Ouvéa include the following taxa: Minysynchiropus laddi, Myripristis
amaenus, Schismorhynchus labialis, Brotula townsendi, Centropyge nigriocellus, and
Centropyge loriculus.
Two species, Alticus sertatus and Neoglyphidodon carlsoni, were previously
thought to be Fijian endemics. The presence of Alticus sertatus at Ouvéa provides the
first record of this species west of the Fiji Islands. An undescribed species of Alticus
occurs at New Caledonia, but was not collected at Ouvéa. Likewise, A/ticus sertatus has
not been collected from New Caledonia. The occurrence of Neoglyphidodon carlsoni at
Ouvéa is the first record outside of Fijian waters.
A comparison of the diversity of the major families within several areas of the
southwestern Pacific is given in Table 2. An interesting analogy with the Chesterfield
archipelago (LeBorgne et al., 1994) and Rotuma (Zug et al., 1989) is the low number or
lack of Abudefduf spp., Neopomacentrus spp., Clupeidae and Siganidae (Table 2). A
preliminary list of fishes from Osprey Reef, off the Great Barrier Reef, based on the fish
collection at the Australian Museum (Leis, pers. communication) entirely lacks these
genera and families. Similarly, Norfolk, Lord Howe and Kermadec islands also have few
or no members of these taxa, but have five species of Abudefduf. A survey of Elizabeth
5
and Middleton Reefs (Hutchings, 1988), located further south in the Coral Sea, indicates
the presence of 3 species of Abudefduf, one species of Siganidae, but no Neopomacentrus
or Clupeidae. These genera and families are well represented elsewhere in the Coral Sea
and New Caledonia (Table 2). At Ouvéa, the lack of suitable habitat is not a likely reason
for the low number of these species, except possibly for the Clupeidae. Indeed, the
habitats where these species are found in New Caledonia are apparently present at Ouvéa
(Kulbicki et al., 1993a). The early life history traits (type of egg, length of larval life,
shape of the larvae, size of the larvae at recruitment on reefs) of Abudefduf and
Neopomacentrus (Table 3) do not differ significantly from those of other Pomacentridae
that are present at Ouvéa, Rotuma, the Chesterfield Islands, or Osprey Reef. Victor
(1991), in a review on settlement strategies and biogeography, noted that duration of the
larval stage rarely accounted for the geographic range of a species. The short distance
between Ouvéa and the main island of New Caledonia (60 km) and the direction of the
major surface currents (Kulbicki et al., 1993a; Kulbicki, 1995) would not seem to present
a major obstacle to the colonization of Ouvéa by species from New Caledonia (figure 3).
Fig. 3. Major currents in the vicinity of Ouvea (from Kulbicki et al., 1993a).
One possible reason for the lack of certain species at Ouvéa could be related to its being
a low island and New Caledonia a high island. Causal factors leading to the absence of
certain taxa at Ouvéa remain speculative.
ACKNOWLEDGEMENTS
We wish to thank the following persons who have helped in the collection or
identification of specimens during the preparation of this checklist: G. Allen, G.
Bargibant, B. Carlson, P. Dalzell, A.C. Gill, B. Hutchins, H. Larson, J. Leis, J.L. Menou,
R. Mooi, G. Mou Tham, S. Poss, J.E. Randall, J. Rivaton, B. Sérét, D.G. Smith. W-F.
Smith-Vaniz, P. Tirard, R. Winterbottom, and the crews of the RV ALIS, DAWA and
DAR MAD. We thank V.G. Springer, A.C. Gill, and K.E. Carpenter for constructive
comments on earlier versions of the manuscript.
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Table 1. List of the shorefish taxa known from Ouvéa atoll. Taxa which were
not previously known from New Caledonia are marked by * and are in
bold typeface. Families are ordered according to Eschmeyer (1990).The
letters used in the column "Method" mean the following:
V: visual census _R: rotenone
Scientific name
GINGLYMOSTOMATIDAE
Nebrius ferrugineus
CARCHARHINIDAE
Carcharhinus albimarginatus
Carcharhinus amblyrhinchos
Carcharhinus melanopterus
Galeocerdo cuvier
Triaenodon obesus
DASY ATIDIDAE
Dasyatis kuhlii
MORINGUIDAE
Moringua species
CHLOPSIDAE
Kaupichthys species
*Kaupichthys atronasus
Kaupichthys hyoproroides
MURAENIDAE
*Anarchias cantonensis
Echidna polyzona
*Echidna unicolor
Gymnothorax buroensis
Gymnothorax chilospilus
Gymnothorax eurostus
Gymnothorax fuscomaculatus
Gymnothorax javanicus
Gymnothorax margaritophorus
*Gymnothorax marshallensis
Gymnothorax melatremus
Gymnothorax pindae
Gymnothorax rueppelliae
Gymnothorax thyrsoides
Gymnothorax zonipectis
Uropterygius fuscoguttatus
*Uropterygius makatei
OPHICHTHIDAE
Leiuranus semicinctus
*Muraenichthys gymnotus
Muraenichthys sp.
*Schismorhynchus labialis
*Schultzidia johnstonensis
CONGRIDAE
Conger cinereus
Heteroconger hassi
CLUPEIDAE
L: line fishing
A: Allen (1975 and pers. comm.)
Author and date
(Lesson, 1830)
(Ruppell, 1837)
(Bleeker, 1856)
(Quoy & Gaimard, 1824)
(Peron & LeSueur, | 822)
(Riippell, 1837)
(Miller & Henle, 1841)
Schultz, 1953
(Stromann, 1896)
(Schultz, 1943)
(Richardson, 1844)
Schultz, 1953
(Bleeker, 1857)
Bleeker, 1865
(Abbott, 1861)
(Schultz, 1953)
(Bleeker, 1859)
Bleeker, 1865
(Schultz, 1953)
(Schultz, 1953)
Smith, 1962
(McClelland, 1845)
(Richardson, 1845)
Seale, 1906
Schultz, 1953
Gosline, 1958
(Lay & Benett, 1839)
Bleeker, 1850
(Seale, 1917)
(Schultz & Woods, 1949)
(Riippell, 1828)
(Klausewitz & Eibl-Eibesfeldt, 1959)
O:1979ORSTOM cruise
Method
WAL
O
AAAAAAA AA << AADAAAA DA
AA OA A
AD”
i
Herklotsichthys quadrimaculatus
Spratelloides delicatulus
CHANIDAE
Chanos chanos
SYNODONTIDAE
Saurida gracilis
Synodus binotatus
Synodus dermatogenys
Synodus jaculum
Synodus hoshinonis
Synodus variegatus
OPHIDIIDAE
Brotula multibarbata
*Brotula townsendi
BY THITIDAE
Brosmophyciops pautzkei
*Dinematichthys randall
*Dinematichthys riukiuensis
ANTENNARIIDAE
Antennarius coccineus
*Antennarius duescus
Antennarius nummifer
GOBIESOCIDAE
Diademichthys lineatus
Discotrema crinophila
*Pherallodichthys species
*Pherallodus species
ATHERINIDAE
*Atherinomorus duodecimalis
Atherinomorus lacunosus
*Atherion elymus
Hypoatherina barnesi
HOLOCENTRIDAE
Myripristis amaena
Myripristis berndti
Myripristis kuntee
Myripristis murdjan ?
Myripristis pralinia
Myripristis violacea
Neoniphon argenteus
Neoniphon opercularis
Neoniphon sammara
Plectrypops lima
Sargocentron caudimaculatum
Sargocentron diadema
Sargocentron melanospilos
Sargocentron punctatissimum
Sargocentron rubrum
Sargocentron spiniferum
Sargocentron tiere
Sargocentron violaceum
AULOSTOMIDAE
Aulostomus chinensis
FISTULARIDAE
(Riippell, 1837)
(Bennett, 1831)
(Forsskal, 1775)
(Quoy & Gaimard, 1824)
Schultz, 1953
Fowler, 1912
Russell & Cressey, 1979
Tanaka, 1917
(Lacépéde, 1803)
Temminck & Schlegel, 1846
Fowler, 1900
Schultz, 1960
Machida, 1994
Aoyagi, 1952
(Lesson, 1831)
Snyder, 1904
(Cuvier, 1817)
(Sauvage, 1883)
Briggs, 1976
(Valenciennes, 1835)
(Schneider, 1 801)
Jordan & Starks,1901
Schultz, 1953
(Castelnau, 1873)
Jordan & Evermann, 1903
Cuvier, 1831
(Forsskal, 1775)
Cuvier, 1829
Bleeker, 1851
(Valenciennes, 1831)
(Valenciennes, 1831)
(Forsskal, 1775)
(Valenciennes, 1831)
(Riippell, 1835)
(Lacépéde, 1801)
(Bleeker, 1858)
(Cuvier, 1829)
(Forsskal, 1775)
(Forsskal, 1775)
(Cuvier, 1829)
(Bleeker, 1853)
(Linnaeus, | 758)
VR
VR
VRO
Zep eof maze)
v2)
Fistularia commersonii
CENTRICIDAE
Aeoliscus strigatus
SYNGNATHIDAE
Corythoichthys amplexus
Corythoichthys nigripectus
Corythoichthys schultzi
Doryrhamphus dactyliophorus
Doryrhamphus excisus excisus
*Phoxocampus diacanthus
SCORPAENIDAE
Dendrochirus brachypterus
Dendrochirus species
Pterois antennata
Pterois radiata
Scorpaena species
Scorpaenodes albaiensis
*Scorpaenodes corallinus
*Scorpaenodes hirsutus
Scorpaenodes kelloggi
Scorpaenodes parvipinnis
Scorpaenodes scaber
Scorpaenopsis species
Scorpaenopsis gibbosa
Scorpaenopsis neglecta
Sebastapistes species
Sebastapistes cyanostigma
*Sebastapistes mauritiana
Sebastapistes strongia
Sebastapistes tinckhami
Setarches species
CARACANTHIDAE
Caracanthus maculatus
Caracanthus unipinna
APLOACTINIDAE
*Neoaploactis tridorsalis
SERRANIDAE
Anyperodon leucogrammicus
Aporops bilinearis
Belonoperca chabanaudi
Cephalopholis argus
Cephalopholis miniata
Cephalopholis sonnerati
Cephalopholis urodeta
Epinephelus caeruleopunctatus
Epinephelus coioides
Epinephelus cyanopodus
Epinephelus fasciatus
Epinephelus hexagonatus
Epinephelus macrospilos
Epinephelus maculatus
Epinephelus merra
Epinephelus polyphekadion
Epinephelus rivulatus
Riippell, 1838
(Ginther, 1860)
Dawson & Randall,1975
Herald, 1953
Herald, 1953
(Bleeker, 1853)
Kaup, 1856
(Schultz, 1943)
(Cuvier, 1829)
(Bloch, 1787)
Cuvier, 1829
(Evermann & Seale,1907)
Smith, 1957
(Smith, 1957)
(Jenkins, 1903)
(Garrett, 1863)
(Ramsay & Ogilby,1886)
(Schneider, 1801)
(Temminck & Schlegel, 1844)
(Bleeker, 1856)
(Cuvier, 1829)
(Cuvier, 1829)
(Fowler, 1946)
(Gray,1831)
(Gray,1831)
Eschmeyer & Allen, 1978
(Valenciennes, 1828)
Schultz, 1943
Fowler & Bean, 1930
Bloch and Schneider, 1801
(Forsskal, 1775)
(Valenciennes, | 828)
(Schneider, 1801)
(Bloch, 1790)
(Hamilton, 1822)
(Richardson, 1846)
(Forrskal, 1775)
(Forster, 1801)
(Bleeker, 1855)
(Bloch, 1790)
Bloch, 1793
(Bleeker, 1849)
(Valenciennes, 1830)
<
AAAAAAADAAAAAAAAAAA << AAAAAA
Ay
wn
1oe)
14
Epinephelus tauvina
Gracila albomarginata
Grammistes sexlineatus
Liopropoma susumi
Liopropoma tonstrinum
Luzonichthys waitei
Luzonichthys williamsi
Plectranthias longimanus
*Plectranthias nanus
Plectranthias winniensis
Plectropomus laevis
Plectropomus leopardus
Pseudanthias ventralis ventralis
Pseudanthias hypselosoma
Pseudanthias lori
Pseudanthias pascalus
Pseudanthias pictilis
*Pseudanthias rubrizonatus
Pseudanthias squamipinnis
Pseudogramma polyacantha
Serranocirrhitus latus
Variola louti
PSEUDOCHROMIDAE
Cypho purpurascens
Pseudochromis species
Pseudochromis cyanotaenia
Pseudochromis jamesi
Pseudoplesiops species
Pseudoplesiops howensis
Pseudoplesiops multisquamatus
Pseudoplesiops rosae
PLESIOPIDAE
Plesiops coeruleolineatus
ACANTHOCLINIDAE
Belonepterygion fasciolatum
KUHLIIDAE
Kuhlia mugil
PRIACANTHIDAE
Heteropriacanthus cruentatus
Priacanthus hamrur
APOGONIDAE
Apogon species
Apogon angustatus
Apogon apogonides
Apogon aureus
*Apogon caudicinctus
Apogon coccineus
Apogon cyanosoma
*Apogon diversus
Apogon doderleini
Apogon doryssa
Apogon erythrinus
Apogon exostigma
Apogon fraenatus
(Forsskal, 1775)
(Fowler & Bean, 1930)
(Thiinberg, 1792)
(Jordan & Seale, 1906)
Randall & Taylor,1988
(Fowler, 1931)
Randall & McCosker,1992
(Weber, 1913)
Randall, 1980
(Tyler, 1966)
(Lacépéde, 1801)
(Lacépéde, 1802)
(Randall, 1979)
Bleeker, 1878
(Lubbock & Randall, 1976)
(Jordan & Tanaka, 1927)
(Randall &Allen, 1978)
(Randall, 1983)
(Peters, 1855)
(Bleeker, 1856)
Watanabe, 1949
(Forsskal, 1775)
(De Vis,1884)
Bleeker, 1857
Schultz, 1943
Allen,1987
Allen, 1987
Schultz, 1943
Riippell, 1835
(Ogilby, 1889)
(Bloch & Schneider, 1 801)
(Lacépéde, 1801)
(Forsskal, 1775)
(Smith & Radcliffe, 1911)
(Bleeker, 1856)
(Lacépéde, 1802)
Randall & Smith, 1988
Rtippell, 1838
Bleeker, 1853
(Smith & Radcliffe, 1911)
Jordan & Snyder, 1901
(Jordan & Seale, 1906)
Snyder, 1904
(Jordan & Starks, 1906)
Valenciennes 1832
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Apogon fuscus
Apogon kallopterus
Apogon nigrofasciatus
Apogon novemfasciatus
Apogon trimaculatus
Apogonichthys ocellatus
Cheilodipterus macrodon
Cheilodipterus quinquelineatus
Fowleria abocellata
Fowleria marmorata
Gymnapogon urospilotus
Pseudamiops gracillicauda
Rhabdamia cypselurus
Rhabdamia gracilis
Siphamia species
SILAGINIDAE
Sillago species
MALACANTHIDAE
Malacanthus brevirostris
Malacanthus latovittatus
ECHENEIDAE
Echeneis naucrates
CARANGIDAE
Alectes indicus
Carangoides chrysophrys
Carangoides ferdau
Carangoides fulvoguttatus
Caranx lugubris
Caranx melampygus
Caranx sexfasciatus
Decapterus russelli
Elagatis bipinnulatus
Gnathanodon speciosus
Scomberoides tol
Trachinotus bailloni
Trachinotus blochii
LUTJANIDAE
Aphareus furca
Aprion virescens
Lutjanus argentimaculatus
Lutjanus bohar
Lutjanus fulviflamma
Lutjanus fulvus
Lutjanus gibbus
Lutjanus kasmira
Lutjanus lutjanus
Lutjanus quinquelineatus
Lutjanus rivulatus
Lutjanus russelli
Lutjanus vitta
Macolor niger
CAESIONIDAE
Caesio caerulaurea
Caesio cuning
Quoy & Gaimard, 1824
Bleeker, 1856
Lachner, 1953
Cuvier, 1828
Cuvier, 1828
(Weber, 1913)
(Lacépéde, 1802)
Cuvier, 1828
Goren & Karplus, 1980
Alleyne & Macleay, 1877
Lachner, 1953
(Lachner, 1953)
Weber, 1909
(Bleeker, 1856)
(Guichenot, 1848)
(Lacépéde, 1801)
Linnaeus, 1758
(Riippell, 1830)
(Cuvier, 1833)
(Forsskal, 1775)
(Forrskal, 1775)
Poey, 1860
Cuvier, 1833
Quoy & Gaimard, 1824
(Riippell, 1830)
(Quoy & Gaimard, 1825)
(Forsskal, 1775)
(Cuvier, 1832)
(Lacépéde, 1801)
(Lacépéde, 1801)
(Lacépéde, 1802)
Valenciennes, 1830
(Forsskal, 1775)
(Forsskal, 1775)
(Forsskal, 1775)
(Bloch & Schneider, 1801)
(Forsskal, 1775)
(Forsskal, 1775)
Bloch, 1790
(Bloch, 1790)
(Cuvier, 1828)
(Bleeker, 1849)
(Quoy & Gaimard, 1824)
(Forsskal, 1775)
Lacépéde, 1801
(Bloch, 1791)
16
Caesio teres
*Gymmnocaesio gymnoptera
*Pterocaesio chrysozona
Pterocaesio digramma
Pterocaesio pisang
*Pterocaesio tessellata
Pterocaesio tile
Pterocaesio trilineata
HAEMULIDAE
Diagramma pictum
Plectorhinchus chaetodonoides
Plectorhinchus goldmanni
Plectorhinchus obscurum
Plectorhinchus picus
LETHRINIDAE
Gnathodentex aurolineatus
Gymnocranius euanus
Gymnocranius grandoculis
Gymnocranius species
Lethrinus atkinsoni
Lethrinus genivittatus
Lethrinus harak
Lethrinus lentjan
Lethrinus miniatus
Lethrinus nebulosus
Lethrinus obsoletus
Lethrinus olivaceus
Lethrinus rubrioperculatus
*Lethrinus species
Lethrinus variegatus
Lethrinus xanthochilus
Monotaxis grandoculis
NEMIPTERIDAE
Scolopsis bilineatus
Scolopsis trilineatus
MULLIDAE
Mutlloides flavolineatus
Mulloides vanicolensis
Parupeneus barberinoides
Parupeneus barberinus
Parupeneus bifasciatus
Parupeneus ciliatus
Parupeneus cyclostomus
Parupeneus heptacanthus
Parupeneus indicus
Parupeneus multifasciatus
Parupeneus pleurostigma
Parupeneus spilurus
Upeneus species (barbillon blanc)
Upeneus species (barbillon jaune)
Upeneus tragula
PEMPHERIDAE
Parapriacanthus ransonneti
Pempheris oualensis
Seale, 1906
(Bleeker, 1856)
(Cuvier, 1830)
(Bleeker, 1865)
(Bleeker, 1853)
Carpenter, 1987
(Cuvier, 1830)
Carpenter, 1987
(Thiinberg, 1792)
Lacépéde, 1800
(Bleeker, 1853)
(Giinther, 1871)
(Cuvier, 1830)
(Lacépéde, 1802)
Giinther, 1879
(Valenciennes, 1830)
Seale, 1909
Valenciennes, 1830
(Forsskal, 1775)
(Lacépéde, 1802)
(Bloch & Schneider, 1801)
(Forsskal, 1775)
(Forsskal, 1775)
Valenciennes 1830
~ Sato, 1978
Valenciennes, 1830
Klunzinger, 1870
(Forsskal, 1775)
(Bloch, 1793)
Kner, 1868
(Lacépéde, 1801)
(Valenciennes, 1831)
(Lacépéde, 1801)
(Lacépéde, 1801)
(Lacépéde, 1801)
(Lacépéde, 1801)
(Lacépéde, 1801)
(Lacépéde, 1801)
(Shaw, 1803)
(Quoy & Gaimard, 1825)
(Bennett, 1830)
(Bleeker, 1854)
Richardson, 1846
Steindachner, 1870
Cuvier, 1831
<< A7A << WAR
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Pempheris swenkii
KYPHOSIDAE
Kyphosus vaigiensis
CHAETODONTIDAE
Chaetodon auriga
Chaetodon baronessa
Chaetodon benetti
Chaetodon citrinellus
Chaetodon ephippium
Chaetodon flavirostris
Chaetodon kleinii
Chaetodon lineolatus
Chaetodon lunula
Chaetodon melanotus
Chaetodon mertensii
Chaetodon ornatissimus
Chaetodon pelewensis
Chaetodon plebeius
Chaetodon rafflesi
Chaetodon reticulatus
Chaetodon speculum
Chaetodon trifascialis
Chaetodon trifasciatus
Chaetodon ulietensis
Chaetodon unimaculatus
Chaetodon vagabundus
Coradion altivelis
Forcipiger flavissimus
Forcipiger longirostris
Hemitaurichthys polylepis
Heniochus acuminatus
Heniochus chrysostomus
Heniochus monoceros
Heniochus singularius
Heniochus varius
POMACANTHIDAE
Centropyge bicolor
Centropyge bispinosus
Centropyge flavissimus
Centropyge heraldi
*Centropyge loriculus
Centropyge multifasciatus
Centropyge nigriocellus
Centropyge tibicen
Centropyge vrolicki
Pomacanthus imperator
Pomacanthus semicirculatus
Pomacanthus sexstriatus
Pygoplites diacanthus
POMACENTRIDAE
Abudefduf sexfasciatus
Abudefduf vaigiensis
Amblyglyphidodon aureus
Amblyglyphidodon leucogaster
Bleeker, 1855
(Quoy & Gaimard, 1825)
Forsskal, 1775
Cuvier, 1831
Cuvier, 1831
Cuvier, 1831
Cuvier, 1831
Giinther, 1873
Bloch, 1790
Cuvier, 1831
(Lacépéde, 1803)
Bloch & Schneider, 1801
Cuvier, 1831
Cuvier, 1831
Kner, 1868
Cuvier, 1831
Bennett, 1830
Cuvier, 1831
Cuvier, 1831
Quoy & Gaimard, 1824
Park, 1797
Cuvier, 1831
Bloch, 1787
Linnaeus, 1758
McCulloch, 1916
Jordan & McGregor, 1898
(Broussonet, 1782)
(Bleeker, 1857)
(Linnaeus, 1758)
Cuvier, 1831
Cuvier, 1831
Smith & Radcliffe, 1911
(Cuvier, 1829)
(Bloch, 1787)
(Giinther, 1860)
(Cuvier, 1831)
Woods & Schultz, 1953
(Giinther, 1874)
(Smith & Radcliffe, 1911)
Woods & Schultz, 1953
(Cuvier, 1831)
(Bleeker, 1853)
(Bloch, 1787)
(Cuvier, 1831)
(Cuvier, 1831)
(Boddaert, 1772)
(Lacépéde, 1802)
(Quoy & Gaimard, 1825)
(Cuvier, 1830)
(Bleeker, 1847)
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Amphiprion akindynos
Amphiprion clarkii
Amphiprion melanopus
Amphiprion perideraion
Chromis acares
Chromis agilis
Chromis amboinensis
Chromis analis
Chromis atripectoralis
Chromis atripes
Chromis chrysura
Chromis flavomaculata
Chromis fumea
Chromis iomelas
Chromis lepidolepis
Chromis margaritifer
*Chromis cf nitida
Chromis retrofasciata
Chromis ternatensis
Chromis vanderbilti
Chromis viridis
Chromis weberi
Chromis xanthochira
Chromis xanthura
Chromis sp.
Chrysiptera biocellata
Chrysiptera leucopoma
Chrysiptera rex
Chrysiptera rollandi
Chrysiptera starki
Chrysiptera taupou
Dascyllus aruanus
Dascyllus reticulatus
Dascyllus trimaculatus
Lepidozygus tapeinosoma
*Neoglyphidodon carlsoni
Neopomacentrus cyanomos
Neopomacentrus violascens
Plectroglyphidodon dickii
Plectroglyphidodon imparipennis
Plectroglyphidodon johnstonianus
Plectroglyphidodon lacrymatus
Plectroglyphidodon leucozonus
*Pomacentrus adelus
Pomacentrus amboinensis
Pomacentrus bankanensis
Pomacentrus brachialis
Pomacentrus chrysurus
Pomacentrus coelestis
Pomacentrus lepidogenys
Pomacentrus molluccensis
Pomacentrus nagasakiensis
Pomacentrus pavo
Pomacentrus philippinus
Allen, 1972
(Bennett, 1830)
Bleeker, 1852
Bleeker, 1855
Randall & Swerdloff, 1973
Smith 1960
(Bleeker, 1873)
(Cuvier, 1830)
Welander & Schultz, 1951
Fowler & Bean, 1928
(Bliss, 1883)
Kamohara, 1960
(Tanaka, 1917)
Jordan & Seale, 1906
Bleeker, 1877
Fowler, 1946
(Whitley, 1928)
Weber, 1913
(Bleeker, 1856)
(Fowler, 1941)
(Cuvier, 1830)
Fowler & Bean, 1928
(Bleeker, 1851)
(Bleeker, 1854)
(Quoy & Gaimard, 1824)
(Lesson, 1830)
(Snyder, 1909)
(Whitley, 1961)
(Allen, 1973)
(Jordan & Seale, 1906)
(Linneaus, 1758)
(Richardson, 1846)
(Riippell, 1828)
(Bleeker, 1856)
(Allen, 1975)
(Bleeker, 1856)
(Bleeker, 1848)
(Lienard, 1839)
(Vaillant & Sauvage, 1875)
Fowler & Ball, 1924
(Quoy & Gaimard, 1824)
(Bleeker, 1859)
Allen, 1991
Bleeker, 1868
Bleeker, 1853
Cuvier, 1830
Cuvier, 1830
Jordan & Starks, 1901
Fowler & Ball, 1928
Bleeker, 1853
Tanaka, 1917
(Bloch, 1787)
Evermann & Seale, 1907
Pomacentrus vaiuli
Pomachromis richardsoni
Stegastes albifasciatus
*Stegastes cf apicalis
Stegastes fasciolatus
Stegastes gascoynel
Stegastes nigricans
CIRRHITIDAE
Amblycirrhitus bimacula
Cirrhitichthys falco
Cyprinocirrhites polyactis
Paracirrhites arcatus
Paracirrhites forsteri
Paracirrhites hemistictus
OPISTOGNATHIDAE
Opistognathus new species
LABRIDAE
Anampses caeruleopunctatus
Anampses geographicus
Anampses neoguinaicus
Anampses twistii
Bodianus anthioides
Bodianus axillaris
Bodianus bilunulatus ?
Bodianus diana
Bodianus loxozonus
Bodianus perditio
Cheilinus bimaculatus
Cheilinus chlorourus
Cheilinus digrammus
Cheilinus trilobatus
Cheilinus undulatus
Cheilinus unifasciatus
Cheilo inermis
Cirrhilabrus laboutei
Cirrhilabrus punctatus
*Cirrhilabrus species
Coris aygula
Coris dorsomacula
Coris gaimard
Coris pictoides
Coris schroederi
Epibulus insidiator
Gomphosus varius
Halichoeres biocellatus
Halichoeres chrysus
Halichoeres hortulanus
Halichoeres margaritaceus
Halichoeres marginatus
Halichoeres melanurus
Halichoeres miniatus
Halichoeres nebulosus
Halichoeres prosopeion
Halichoeres trimaculatus
Jordan & Seale, 1906
(Snyder, 1909)
(Schlegel & Miller, 1839-44)
(De Vis, 1885)
(Ogilby, 1889)
(Whitley, 1964)
(Lacépéde, 1802)
Jenkins, 1903
Randall, 1963
(Bleeker, 1875)
(Cuvier, 1829)
(Schneider, 1801)
(Giinther, 1874)
Smith-Vaniz, in prep.
Riippell, 1829
Valenciennes, 1840
Bleeker, 1878
Bleeker, 1856
(Bennett, 1830)
(Bennett, 1831)
(Lacépéde, 1801)
(Lacépéde, 1801)
(Snyder, 1908)
(Quoy & Gaimard, 1834)
Valenciennes, 1840
(Bloch, 1791)
(Lacépéde, 1801)
Lacépéde, 1801
Riippell, 1835
Streets, 1877
(Forsskal, 1775)
Randall & Lubbock, 1982
Randall & Kuiter, 1989
Lacépéde, 1801
Fowler, 1908
(Quoy & Gaimard, 1824)
Randall & Kutter, 1982
(Bleeker, 1858)
(Pallas, 1770)
Lacépéde, 1801
Schultz, 1960
Randall, 1981
(Lacépéde, 1801)
(Valenciennes, 1839)
Riippell, 1835
(Bleeker, 1851)
(Valenciennes, 1839)
(Valenciennes, 1839)
(Bleeker, 1853)
(Quoy & Gaimard, 1834)
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Hemigymnus fasciatus
Hemigymnus melapterus
*Hologymnosus annulatus
Hologymnosus doliatus
Labrichthys unilineatus
Labroides bicolor
Labroides dimidiatus
Labroides pectoralis
Labropsis australis
Labropsis xanthonota
Macropharyngodon meleagris
Macropharyngodon negrosensis
Novaculichthys taeniorus
Pseudocheilinus evanidus
Pseudocheilinus hexataenia
Pseudocheilinus octotaenia
Pseudocheilinus species
Pseudocoris yamashiroi
*Pseudodax moluccanus
Pseudojuloides cerasinus
Stethojulis bandanensis
Stethojulis interrupta
Suezichthys gracilis
Thalassoma amblycephalum
Thalassoma hardwicke
Thalassoma jansenii
Thalassoma lunare
Thalassoma lutescens
Thalassoma purpureum
Thalassoma quinquevittatum
Thalassoma trilobatum
Wetmorella nigropunctata
Xyrichthys pavo
SCARIDAE
Bolbometopon muricatum
Cetoscarus bicolor
Hipposcarus longiceps
Scarus altipinnis
Scarus chameleon
Scarus flavipectoralis
Scarus forsteni
Scarus frenatus
Scarus ghobban
Scarus globiceps
Scarus longipinnis
Scarus microrhinos
Scarus niger
Scarus oviceps
Scarus psittacus
Scarus rivulatus
Scarus rubroviolaceus
Scarus schlegeli
Scarus sordidus
Scarus spinus
(Bloch, 1792)
(Bloch, 1791)
(Lacépéde, 1801)
(Lacépéde, 1801)
(Guichenot, 1847)
Fowler & Bean, 1928
(Valenciennes, 1839)
Randall & Springer, 1975
Randall, 1981
Randall, 1981
(Valenciennes, 1839)
Herre, 1932
(Lacépéde, 1801)
Jordan & Evermann, 1903
(Bleeker, 1857)
Jenkins, 1900
(Schmidt, 1930)
(Valenciennes, 1839)
(Snyder, 1904)
(Bleeker, 1851)
(Bleeker, 1851)
(Steindachner & Doderlein, 1887)
(Bleeker, 1856)
(Bennett, 1828)
(Bleeker, 1856)
(Linnaeus, 1758)
(Lay & Bennett, 1839)
(Forsskal, 1775)
(Lay & Bennett, 1839)
(Lacépéde, 1801)
(Seale, 1901)
Valenciennes, 1840
(Valenciennes, 1840)
(Riippell, 1829)
(Valenciennes, 1840)
(Steindachner, 1879)
Choat & Randall, 1986
Schultz, 1958
(Bleeker, 1861)
Lacépéde, 1802
Forsskal, 1775
Valenciennes, 1840
Randall & Choat, 1980
Bleeker, 1854
Forsskal, 1775
Valenciennes, 1840
Forsskal, 1775
Valenciennes, 1840
Bleeker, 1847
(Bleeker, 1861)
Forsskal, 1775
(Kner, 1868)
aT SW Ch RR eR ay
CREEDIIDAE
*Chalixodytes chameleontoculis
*Chalixodytes tauensis
Limnichthys donaldsoni
PINGUIPEDIDAE
Parapercis clathrata
Parapercis cylindrica
Parapercis hexophtalma
Parapercis millepunctata
Parapercis multiplicata
Parapercis schauinslandi
TRIPTERYGIIDAE
Ceratobregma helenae
Ceratobregma striata
Enneapterygius elegans
Enneapterygius flavoccipitis
Enneapterygius hemimelas
Enneapterygius nanus
Enneapterygius niger
Enneapterygius rufopileus
Enneapterygius tutuilae
Helcogramma cf ellioti
Norfolkia squamiceps
Norfolkia thomasi
Springerichthys kulbickii
BLENNIIDAE
*Alticus sertatus
Aspidontus dussumieri
Aspidontus taeniatus
Blenniella chrysospilos
Cirripectes castaneus
Cirripectes stigmaticus
Ecsenius bicolor
Ecsenius fourmanoiri
Ecsenius midas
Ecsenius nalolo
Entomacrodus caudofasciatus
*Entomacrodus decussatus
Entomacrodus striatus
Exalias brevis
Istiblennius edentulus
Meiacanthus atrodorsalis
Petroscirtes mitratus
Plagiotremus laudandus
Plagiotremus rhinorhynchos
Plagiotremus tapeinosoma
CALLIONY MIDAE
Diplogrammus goramensis
*Minysynchiropus laddi
Synchiropus morrisoni
Synchiropus ocellatus
ELEOTRIDIDAE
Calumia godeffroyi
GOBIIDAE
Smith, 1956
Schultz, 1943
Schultz, 1960
Ogilby, 1911
(Bloch, 1792)
(Cuvier, 1829)
(Giinther, 1860)
Randall, 1984
(Steindachner, 1900)
Holleman, 1987
Fricke, 1991
(Peters, 1877)
Shen & Wu, 1994
(Kner & Steindachner, 1866)
(Schultz, 1960)
Fricke, 1994
(Waite, 1904)
Jordan & Seale, 1906
(Herre, 1944)
(McCulloch & Waite, 1916)
Whitley, 1964
Fricke & Randall, 1994
(Garman, 1903)
(Valenciennes, 1836)
Quoy & Gaimard, 1834
(Bleeker, 1857)
(Valenciennes, 1836)
Strasburg & Schultz, 1953
(Day, 1888)
Springer, 1972
Starck, 1969
Smith, 1959
(Regan, 1909)
(Bleeker, 1858)
(Quoy & Gaimard, 1836)
(Kner, 1868)
(Bloch, 1801)
(Giinther, 1877)
Riippell, 1830
(Whitley, 1961)
(Bleeker, 1852)
(Bleeker, 1857)
(Bleeker, 1858)
(Schultz, 1960)
Schultz, 1960
(Pallas, 1770)
(Giinther, 1 877)
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Amblyeleotris steinitzi
Amblygobius phalaena
Callogobius species
Callogobius maculipinnis
Callogobius sclateri
Eviota species |
Eviota species 2
Eviota afelei
Eviota albolineata
Eviota cometa
*Eviota fasciola
*Eviota latifasciata
Eviota melasma
Eviota monostigma
Eviota prasinia
Eviota prasites
*Eviota pseudostigma
Eviota sparsa
Eviota zebrina
*Eviota zonura
Coryphopterus duospilos
Coryphopterus neophytus
Gnatholepis cauerensis
Gnatholepis scapulostigma
Gobiodon citrinus
Gobiodon rivulatus
Gobiopsis species
Istigobius decoratus
Istigobius rigilius
Macrodontogobius wilburi
Paragobiodon lacunicolus
Paragobiodon melanosomus
Paragobiodon xanthosomus
Pleurosicya species
Priolepis cincta
*Priolepis compita
Priolepis fallacincta
*Priolepis kappa
Priolepis semidoliatus
*Sueviota lachneri
Trimina 5 species
Trimma caesiura
Trimma okinawae
*Trimma taylori
Trimma tevegae
Trimma unisquamus
Trimmatom eviotops
Trimmatom nanus
Valenciennea puellaris
Valenciennea strigata
KRAEMERIDAE
*Kraemeria samoensis
MICRODESMIDAE
Nemateleotris magnifica
(Klausewitz, 1974)
(Valenciennes, 1837)
(Fowler, 1918)
(Steindachner, 1880)
Jordan & Seale, 1906
Jewett & Lachner, 1983
Jewett & Lachner, 1983
Karnella & Lachner, 1981
Jewett & Lachner, 1983
Lachner & Karnella, 1980
Fourmanoir, 1971
Klunzinger, 1871
Jordan & Seale, 1906
Lachner & Karnella, 1980
Jewett & Lachner, 1983
Lachner & Karnella, 1978
Jordan & Seale, 1906
Hoese & Reader, 1985
(Giinther, 1877)
(Bleeker, 1853)
Herre, 1953
(Riippell, 1838)
(Riippell, 1830)
(Herre, 1927)
(Herre, 1953)
Herre, 1936
(Kendall & Goldsborough, 1911)
(Bleeker, 1852)
(Bleeker, 1852)
(Regan, 1908)
Winterbottom, 1985
Winterbottom & Burridge, 1992
Winterbottom & Burridge, 1992
(Valenciennes, 1837)
Winterbottom & Hoese, 1988
Jordan & Seale, 1906
(Aoyagi, 1949)
Lobel, 1979
Cohen &Davis 1969
(Gosline, 1959)
(Schultz, 1943)
Winterbottom & Emery, 1981
(Tomiyama, 1956)
(Broussonet, 1782)
Steindachner, 1906
Fowler, 1928
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Ptereleotris evides
Ptereleotris hanae
SIGANIDAE
Siganus argenteus
Siganus punctatus
Siganus spinus
ZANCLIDAE
Zanclus cornutus
ACANTHURIDAE
Acanthurus albipectoralis
Acanthurus blochii
Acanthurus dussumieri
Acanthurus guttatus
Acanthurus lineatus
Acanthurus mata
Acanthurus nigricans
Acanthurus nigricauda
Acanthurus nigrofuscus
Acanthurus olivaceus
Acanthurus pyroferus
Acanthurus triostegus
Acanthurus xanthopterus
Ctenochaetus binotatus
Ctenochaetus striatus
Naso annulatus
Naso brachycentron
Naso brevirostris
Naso hexacanthus
Naso lituratus
Naso tuberosus
Naso unicornis
Paracanthurus hepatus
Zebrasoma scopas
Zebrasoma veliferum
SPHYRAENIDAE
Sphyraena barracuda
Sphyraena forsteri
Sphyraena putnamie
SCOMBRIDAE
Euthynnus affinis
Grammatorcynus bicarinatus
Gymnosarda unicolor
Katsuwonus pelamis
Rastrelliger kanagurta
Scomberomorus commerson
BOTHIDAE
Bothus mancus
Bothus pantherinus
PLEURONECTIDAE
Samariscus triocellatus
SOLEIDAE
Soleichthys heterorhinos
BALISTIDAE
Balistapus undulatus
(Jordan & Hubbs, 1925)
(Jordan & Snyder, 1901)
(Quoy & Gaimard, 1825)
(Forster, 1801)
(Linnaeus, 1758)
(Linnaeus, 1758)
Allen & Ayling, 1987
Valenciennes, 1835
Valenciennes, 1835
Forster, 1801
(Linnaeus, 1758)
Cuvier, 1829
(Linnaeus, 1758)
Duncker & Mohr, 1929
(Forsskal, 1775)
Forster, 1801
Kittlitz, 1834
(Linnaeus, 1758)
Valenciennes, 1835
Randall, 1955
(Quoy & Gaimard, 1825)
(Quoy & Gaimard, 1825)
(Valenciennes, 1835)
(Valenciennes, 1835)
(Bleeker, 1855)
(Forster, 1801)
Lacépéde, 1802
(Forsskal, 1775)
(Linnaeus, 1766)
(Cuvier, 1829)
(Bloch, 1797)
(Walbaum, 1792)
Cuvier, 1829
Jordan & Seale, 1905
(Cantor, 1849)
(Quoy & Gaimard, 1824)
(Riippell, 1838)
(Linnaeus, 1775)
(Cuvier, 1817)
(Lacépéde, 1800)
(Broussonet, 1782)
(Riippell, 1828)
Woods, 1966
(Bleeker, 1856)
(Park, 1797)
23
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Balistoides conspicillum
Balistoides viridescens
Melichthys vidua
Pseudobalistes fuscus
Rhinecanthus aculeatus
Rhinecanthus rectangulus
Sufflamen bursa
Sufflamen chrysopterus
Sufflamen fraenatus
MONACANTHIDAE
Amanses scopas
Cantherines dumerili
Oxymonacanthus longirostris
Paraluteres prionurus
Pervagor janthinosoma
Pervagor melanocephalus
OSTRACIIDAE
Ostracion cubicus
Ostracion meleagris
Tetrasoma gibbosus
TETRAODONTIDAE
Arothron hispidus
Arothron meleagris
Arothron nigropunctatus
Canthigaster bennetti
Canthigaster janthinoptera
Canthigaster valentini
Lagocephalus scleratus
DIODONTIDAE
Diodon hystrix
(Bloch & Schneider, 1801)
(Bloch & Schneider, 1801)
(Solander, 1844)
(Bloch & Schneider, 1801)
(Linnaeus, 1758)
(Bloch & Schneider, 1801)
(Bloch & Schneider, 1801)
(Bloch & Schneider, 1801)
(Latreille, 1804)
Cuvier, 1829
(Hollard, 1854)
(Bloch & Schneider, 1801)
(Bleeker, 1851)
(Bleeker, 1854)
(Bleeker, 1853)
Linnaeus, 1758
Shaw, 1796
(Linnaeus, 1758)
(Linnaeus, 1758)
(Lacépéde, 1798)
(Bloch & Schneider, 1801)
(Bleeker, 1854)
(Bleeker, 1855)
(Bleeker, 1853)
(Forster, 1788)
Linnaeus, 1758
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25
Table 2. Shallow water and reef fish species diversity of the major families at Ouvéa and
other areas of the Western Pacific. (Footnote sources are in parentheses.)
Family Ouvéa New Chesterfield South North PNG Norfolk, Fiji Rotuma
Caledonia GBR GBR Lord Howe,
Kermadec
(10) (11) (12) (13) (14) (15) (16) (17)
Acanthuridae WS 33 26 Ds) 36 33 13 23 12
Apogonidae Dy 65 47 34 47 80 10 40 Di
Balistidae 10+6 14+10 9+10 11+11 17+19 17+15 5+10 14+1 1 6+3
+Monacanthidae
Blenniidae 20 43 DY 40 50 63 20 38 32
Caesionidae (1) 10 9 6 4 7 11 2 8 3
Carangidae 13 25 2 32) 45 53 De 28 7
Chaetodontidae 3] 32 23 32 45 44 23 32 14
Clupeidae (2) 2 10 l 5 10 Dy 2 1] 0
Gobiidae 54 84 55 104 153 161 27 na 54
Haemulidae 5 9 3 8 10 18 4 4 0
Holocentridae 18 21 20 1] 25 25 6 DD, 17
Labridae 69 84 73 69 106 93+ 56 64 28
Lethrinidae (3) 17 18 14 9 20 25 5 17 1]
Lutjanidae (4) 14 19 10 14 24 3] 8 20 DD,
Mullidae 15 15 2 7 16 14+ 10 18 9
Muraenidae 17 25 19 23 30 40 17 24 8
Nemipteridae (5) 2 9 py) 7 11 15 | 8 0
Platycephalidae 0 9 10 4 10 late | 2 2
Pomacanthidae 13 15 12 15 24 23 7 16 3
Pomacentridae (6) 58 82 54 69 106 105 35 67 37
Abudefduf 2 6 0 6 6 7 5 5 0
Neopomacentrus 2 6 0 2 6? 7 0 l l
Scaridae 20 26 21 23 DY DY 13 22+ 3
Scorpaenidae 20 23 27 21 26 34 13 17 10
Serranidae (7) 39 47 29 32 88 73 22 42 23
Siganidae (8) 3 9 y) 8 10 13 l 6 l
Syngnathidae (9) 7 29 16 12 18 45 6 32 9
Synodontidae 6 7 7 8 8 12 6 5 4
Tetraodontidae 7 19 10 1] 14 23 9 13 5
Longitude (°E) 167 165 159 154 143 140 165 178 177
Latitude (°S) 20 21 19 22 18 10 30 17 12
Rank in Landmass 6 3 8 ] l yD) 5 4 i
1: Carpenter 1987,1988 2: Whitehead 1985: Whitehead et al., 1988 3: Carpenter and Allen, 1989
4: Allen and Talbot, 1985 5: Russell, 1990 6: Allen, 1991
7: Randall and Heemstra, 1991 8: Woodland, 1990 9: Dawson, 1985
10: Rivaton et al., 1989 11: LeBorgne et al., 1994 12: Russell, 1983: Lowe and Russell, 1990
13: Allen 1989; Randall et al.. 1990, Paxton et al., 1978 14: Kailola, 1987a.b, 1991; Allen and Swainston, 1992
15: Francis, 1993 16: Carlson ms; Blaber et al., 1993 17: Zug et al., 1989
26
Table 3. Characteristics of the early life history traits of Abudefduf and Neopomacentrus
spp. compared to other Pomacentridae.
Genus
Abudefduf
Chromis
Chrysiptera
Dascyllus
Neopomacentrus
Paraglyphidodon
Plectroglyphidodon
Pomacentrus
Stegastes
1 - Brothers et al. 1983
Egg size (mm?)
.308-.450 (3)
.091-.109 (3)
.347-.539 (3)
.093-.181 (3)
.850-.898 (3)
.112-.180 (3)
.210-.804 (3)
112 (3)
2 - Wellington and Victor 1989
Larval duration
(days)
23 (1); 17-20 (2); 22.1-24.2
(3)
20-26 (1): 18-38 (2): 20.3-
28.8 (3)
23-24 (1); 14-24 (2); 17.4-
22.0 (3)
20-28 (1); 16-30 (2): 22.4-
24.2 (3)
24 (1); 16-24 (2):18.2 (3)
14-28 (2): 14.8 (3)
19-33 (2); 24.3 (3)
17-85 (1); 14-27 (2): 19.0-
26.0 (3)
19-39 (2): 32.0 (3)
Size at settlement
(mm)
10.4 (1): 11.2 (2)
8.3 (1): 8-14.4 (2); 8.2-8.9
(3)
10.9 (1); 8.7-11.2 (2): 9.9-
10.9 (3)
7.0 (1): 6.4-10.1 (2):7.0 (3)
ISDE WOES AE 132) (3)
8.6-9.0 (2)
11.9-22.8 (1); 10.6-14.1 (2);
12.2-15.0 (3)
9.5-13.6 (2)
3 - Thresher and Brothers 1989
ATOLL RESEARCH BULLETIN
NO. 445
ON THE ORIGIN OF DRIFT MATERIALS IN THE MARSHALL ISLANDS
BY
D.H.R. SPENNEMANN
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
OCTOBER 1997
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Figure 1 Index map of the Marshall Islands showing the atolls mentioned in the text.
ON THE ORIGIN OF DRIFT MATERIALS IN THE MARSHALL ISLANDS
BY
Dirk H.R. Spennemann)
The oceanic dispersal of plants and animals has been the focus of studies ever since
organized natural history started in the Pacific, and the dispersal of terrestrial by sea
rafting has been given due consideration. The finding of drift materials such as glass floats,
tree trunks and seeds, is a common occurrence on the shores of Pacific Islands, but in most
cases the origin of such material is unknown or at least equivocal. Thus while the principle
of sea rafted dispersal is known and reported at length, there is a need to document those
occasions where positive proof of origin can be furnished.
Recently a piece of pumice with a slab of obsidian (volcanic glass) attached to it was
found on an atoll in the Marshall Islands—a coral atoll group devoid of volcanic materials
(Spennemann & Ambrose 1997). It became necessary to review the archaeological and
historical record of the nature and origin of drift materials washed up on the atolls of the
Marshall Islands (Spennemann 1996). In view of the relevance of this information for
biogeographical studies in general it seems prudent to furnish the salient points in a format
accessible to a wider academic community.
THE MARSHALL ISLANDS
The Marshall Islands, comprising 29 atolls and 5 islands, are located in the north-west
equatorial Pacific, about 3790km west of Honolulu, about 2700km north of Fiji and
1500km east of Ponape. With the exception of the two northwestern atolls, Enewetak and
Ujelang, the Marshall Islands are arranged in two island chains running roughly NNW to
SSE: the western Ralik Chain and the eastern Ratak Chain (figure 1).
The current patterns in the Marshall Islands are complex and material can float in from
both the east and the west. Three current zones can be encountered, which are the south
equatorial current, running from east to west, the equatorial counter current, running from
west to east, and the northern equatorial current running from east to west. During the
northern summer the atolls south of Mile are located within the north equatorial counter
current, which runs against the tradewinds (west to east). In the following northern winter
these atolls are at or near the interface between the north equatorial counter current and the
northern equatorial current (running east to west) (Barnes ef al. 1948). In addition, the El
Nifio effect changes the sea surface temperatures and hence the climatic belts. Further,
typhoons, whose frequencies seem to be running in synchrony with the occurrence of the
El Nifio effect (Spennemann & Marschner 1995), bring material from other destinations to
the Marshalls.
J The Johnstone Centre, Charles Sturt University, PO Box 789, Albury NSW 2640,
Australia.
Manuscript received 15 June 1996; revised 27 June 1997
The first European account of the occurrence of driftwood in the Marshall Islands was
reported by Adalbert von Chamisso who noted that it was seen on Wotje in 1816 on
occasion of the visit by the Russian Exploring Expedition commanded by Otto von
Kotzebue (Chamisso 1910, p. 156). Driftwood is a common occurrence throughout the
atolls of the Marshall Islands (Hager 1886, p. 57), and has been reported from the
shoreline of many atolls, such as Majuro (Spennemann 1992); Arno (Wells 1951, p. 3);
Mile (pers. obs. ); Wotje (Chamisso 1910); Kwajalein (Fosberg 1956: plate 13A); Ebon
(pers. obs.), Nadikdik (pers obs.); Maloelap (pers obs.); and Jaluit (Schneider 1891). It has
also been found in the centre of other islets, such as on Wake (Eneen-Kio) Atoll (Grooch
1936, p. 92; 1938) or Bokak (Taongi) Atoll (Irmer 1895), both in the northern Marshalls.
HISTORIC EVIDENCE FOR THE ORIGIN OF DRIFT MATERIAL
Only a few of the drift items encountered on the shores of the Marshall Islands allow an
accurate identification of their origin. References to these is scattered are the literature.
These are discussed below and summarised in table 1.
Wells reported that driftwood trees arriving from North America (mainly California)
and carried by the Northern Equatorial Current are not uncommon on the atolls of the
Marshall Islands. He encountered a cut fir log measuring 1.5 by 16.5m (5 by 55 feet) and a
trunk of a redwood tree on Arno Atoll (Wells 1951, p. 3). Grooch (1936) reported the
presence of drift logs on Wake and the current author has seen large cut drift logs in
various islands on Majuro, Mile and Nadikdik Atolls. Traditional Marshallese culture has
several references to fir trees and their uses (compiled in Spennemann 1996).
Another report on drift material coming from the east is the case of a rubber dinghy. On
27 May 1943 a Consolidated B-24 ‘Liberator’ bomber crashed into the sea 225m NW of
Palmyra Atoll (crash site approx 8°32'N 164°20'W). The three survivors of the US crew
drifted for 47 days in an inflatable life raft and eventually arrived on Japanese-held
Maloelap Atoll, having floated across the reef into the lagoon (JICPOA 1944).
In a similar case, a small fishing vessel went missing in 1979 off Hana, Maui, Hawaiian
Islands and was eventually found washed up on Bokak Atoll in 1989 (Thomas 1989, p.
33).
Drift voyages by canoes from the west are also documented: Lamotrekese are reported
on Arno (Chamisso 1986, p. 264; Kotzebue 1821, p. II 89), Pingelapese arrived on Jaluit
(Kramer & Nevermann 1938, p. 35) and on different occasions, Yapese drifted to Aur
Atoll (18 century, Chamisso 1986, p. 264) and Kili Island (Hezel 1979, p. 127; entry for
1868, Bark Syringia). In addition canoes from Woleai arrived in the Marshalls (Chamisso
1986; Erdland 1914, p. 315).
Before the introduction of bamboo plants to the Marshalls by the Japanese colonial
administration, sea rafted bamboo was much sought after for use as bamboo containers and
the like (Kramer & Nevermann 1938; Knappe collection Erfurt, unpubl.). The rafted
bamboo came from sources in South East Asia, most likely Indonesia or the Philippines.
3
Table I Known points of origin for drift materials encountered on atolls of the Marshall
Islands
Locality Target Item floated
Apaiang, Kiribati Mile canoe load of people
California, North America Arno, Majuro, Mile, cut fir logs
Nadikdik, Wake
Central Solomons Mile, Wotje canoe hull
Japan boats
Kiribati (general) Ebon, Namorik canoe load of people
Kiribati (general) Mile sailboat hull
Krakatau, Indonesia Ailuk etc. pumice
Lamotrek Atoll, FSM Arno canoe load of people
Maui, Hawaii Bokak skiff
Palmyra Atoll (225 nmNW Maloelap rubber dinghy from crashed B-
Olleee) 24
Philippines (?) bamboo
Pingelap Atoll, FSM Jaluit canoe load of people
Tuluman I., Bismarcks, PNG _—_ Nadikdik piece of pumice/obsidian
Woleai Atoll, FSM Kili canoe load of people
Yap, FSM Kili, Aur canoe load of people
Similarly, following the explosion of Krakatau in 1883 large amounts of pumice were
produced which were washed ashore in the Marshall Islands (Grundemann 1887, p. 442;
Sachet 1955).
Even though the Marshall Islands’ atolls are in the zone of the north equatorial counter
current, the origin of objects from Japan cannot be excluded. There are abundant examples
of Japanese Junks drifting to Siberia, Alaska, Canada, mainland USA, Hawaii, the
Marianas and Guam, Palau and even the Marshall Islands (Kakubayashi 1983).
In addition, material from sources south of the equator has been documented. During a
pedestrian survey on Nadikdik Atoll (5°45' N, 172°10' E) the author found a piece of
pumice with a slab of obsidian attached to it. As the atoll had been completely water
washed with a 12m high storm surge and devastated with a large loss of life during a
typhoon in 1905, it was very likely that the deposition of the pumice occurred after that
time. Quantitative chemical analysis of major elements in the obsidian showed that
Tuluman Island in the Bismarck Archipelago is the most probable source of the material
(Spennemann & Ambrose in press). Tuluman Island emerged from the sea in 1953 and
produced massive amounts of pumice and obsidian in periodic eruptions until around 1957
(Reynolds ef al. 1980).
Washed up canoe hulls are other indicators of drift materials in the Marshall Islands.
The author has seen two canoe hulls which appear to be Solomon Islands canoes. One was
seen on Mile Atoll, the other, a Binabina-style canoe from the central Solomon Islands, on
Wotje Atoll (Spennemann 1996).
GO°N g
50°N oo
oe ae
120°E 150°E 180° 150°W 120°W 90°wW
Figure 2. Map of the Pacific Ocean showing the origin of the drift materials encountered in
the Marshalls. The greyed dot indicates the location of the southern atolls of the
Marshall Islands
I-Kiribati canoes (with or without crew) were often found adrift. A sail boat hull of a
modern Kiribati design drifted ashore in Mile in the late 1980s and has been refurbished
since (own obs.) I-Kiribati canoes were often stranded on the southern Marshalls,
especially Arno and Mile, and these atolls have several genealogical links with the northern
and central atolls of Kiribati. Shipwrecked i-Kiribati crew were picked up by the brig
Mercury south of Ebon in 1858 (Hezel 1979, p. 121). In 1882 other i-Kiribati were found
drifting south of Ebon by the American vessel Northern Light (Hezel 1979, p. 139).
During the 19 century dispersed i-Kiribati were also living on Namorik (1851; Hezel
1979, p. 121; 1868; ibid. 127) and Jaluit (1871; ibid. 129; 1879 ibid. 136). Two Catholic
missionaries together with fourteen Gilbertese left Apaiang Atoll en route to Marakei in
early September 1942. The canoe eventually wrecked on Mile Atoll in the Marshalls
(Richard 1957, p. 401). Even today, i-Kiribati fishermen occasionally drift to the shores of
the southern Marshall Islands.
In addition, there is evidence for internal drift in the Marshall Islands archipelago.
Following the 1905 typhoon that hit the southern atolls of Nadikdik, Mile, Arno and
5
Jaluit, the remains of canoes, wooden bowls, houses and corpses were washed ashore on
Enewetak Atoll (Jeschke 1906). Following a devastating typhoon in 1840 survivors from
Mejit Island were washed ashore on Likiep Atoll (Erdland 1914, p. 18). Both of these
cases of east to west drift are well within the range of expectations given the overall
current pattern.
160 162 164 166 168 170 172 174
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Figure 3. Map of the Marshall Islands showing the direction from where drift materials cam from. Small
letters designate drift within the Marshalls.
IMPLICATIONS
The locations of confirmed origin of sea rafted materials have been plotted in figure 2. It
becomes evident that material from all areas of the Pacific (with the exception—so far—of
South America and Australia proper) has arrived in the Marshall Islands. The distribution
of the origin of sourced materials on the various Marshall Islands atolls has been plotted in
figure 3 which shows that the southern atolls are more favoured in this respect than the
northern locales.
The perusal of current pattern charts provides are too coarse a resolution and does not
allow to assess micro variations, a scale that is required to make useful predictions For
example, a mere perusal of the current charts would not have made likely for example the
dispersal of material from the Solomons or the Bismarck Archipelago.
As this small compilation has shown, there is still a need to systematically compile and
draw on the historic literature and make use other contemporary material and sources.
Coupled with a reassessment of the Holocene sea-level curve this observation may have a
bearing on the interpretation of the distribution of mangrove species in Eastern
Micronesia.
BIBLIOGRAPHY
Barnes, C.A., D.F. Bumpus and J. Lyman (1948) Ocean circulation in the Marshall Islads
area. Transactions of the American Geophysical Union 29(6): 871-876.
Chamisso, A. von (1910) Reise um die Welt mit der Romanzoffschen Endeckungs-
expedition in den Jahren 1815-1818 auf der Brig Rurik, Capitan Otto v. Kotzebue
Zweiter Theil: Bemerkungen und Ansichten. Chamisso's Werke Vierter Theil. Berlin:
G.Hempel.
Chamisso, A. von (1986) A voyage around the world with the Romanzov exploring
expedition in the years 1815-1818 in the Brig Rurick, Captain Otto von Kotzebue.
(translated by H.Kratz). Honolulu: University of Hawaii Press.
Erdland, A. (1914) Die Marschall-Insulaner. Anthropos Ethnologische Bibliothek. Miinster
i.W. Bd. 2, Heft 1.
Fosberg, F.R. (1956) Military Geography of the Northern Marshalls. Engineer Intelligence
Dossier, Strategic Study Marshall, Subfile 19, Analysis of the Natural Environment.
Prepared under the direction of the Chief of Engineers, U.S. Army by the Intelligence
Division Office of the Engineer Headquarters United States Army Forces Far East
with personnel of the United States Geological Survey.
Grooch, W.S. (1936) Skyway to Asia. New York: Longman & Green.
Grundemann, D. (1887) Unser kleinstes Schutzgebiet, die Marschallinseln. Deutsche
Kolonialzeitung 4, 441-444.
Hager, C. (1886) Die Marschall-Inseln. Leipzig.
Hezel, F. X. (1979) Foreign ships in Micronesia. A compendium of ship contacts with the
Caroline and Marshall Islands 1521-1885. Saipan, Mariana Is.: F.J.Hezel & Trust
Territory Historic Preservation Office.
Irmer, G. (1895) Letter Kaiserlicher Landshauptmann fiir das Schutzgebiet der Marschall
Inseln Dr. Irmer to Reichskanzler Fiirst zu Hohenlohe-Schilligsfiirst. Trip report on a
voyage to Bikar and Bokak. Letter dated Jaluit 14 December 1895. in Auswartiges
Amt Kolonial-Abtheilung AIII Acten betreffend Deutsche Siidsee Phosphat
Gesellschaft. Gesellschaften und Vereine 10f N. 4. File 2459 Vol 1. June 1906 to 15
March 1907. National Library of Australia, Microfilm MfM G 8525.
Jeschke, C. (1906) Bericht tiber die Marschall-Inseln. Petermanns Mitteilungen 52, 270-
UT
JICPOA (1944) "Prisoner of War Interrogation Report, 6th Base Force Secret Number
330, Headquarters 6th Base Force July 1943" and 'Prisoner of War Interrogation
Report Annex 23 July 1943" translation of captured Japanese document JICPOA Item
5703 captured at Kwajalein Atoll, Received JCPOA 11 February 1944 contained in
file "Joint Intelligence Centre Pacific Ocean Areas. Translations of Japanese
documents captured Makin—Kwayjalein Atoll-Namur Island—Munda and Tarawa. Also
primary interrogation of Japanese prisoners of war, March 1944. Record series
AWM5S4 file 423/4/40 Part 1. Archives of the Australian War Memorial, Canberra,
Australia.
Kakuyabashi, F. (1983) Japanese drift records and the Sharp hypothesis. Journal of the
Polynesian Society 90, 515-524.
Kotzebue, O. von (1821) A voyage of discovery into the South Sea and Beering’s Straits.
for the purpose of exploring a north-east passage undertaken in the years 1815-1818,
at the expense of His Highness the Chancellor of the Empire, Count Romanzoff in the
ship Rurick, under the command of the Lieutenant in the Russian Imperial Navy, Otto
von Kotzebue. 3 vols. London: Longman, Hurst, Rees, Orme and Brown.
Kramer, A. & H. Nevermann (1938) Ralik-Ratak (Marschall Inseln). Jn: G.Thilenius
(ed.), Ergebnisse der Stidsee-Expedition 1908-1910. II. Ethnographie, B:
Mikronesien. Vol. 11: Hamburg: Friedrichsen & de Gruyter.
Reynolds, M.A., J.G. Best and R.W. Johnson (1980) The 1953-57 eruption of Tuluman
volcano: rhyolitic activity in the northern Bismark Sea. Geological Survey of Papua
New Guinea, Memoir 7
Richard, D. E. (1957) The United States Naval Administration of the Trust Territory of the
Pacific Islands. Vol. 1: The Wartime Military Government Period 1942-1945,
Washington, DC: U.S. General Printing Office.
Sachet, M.-H. (1955) Pumice and other extraneous volcanic material on coral atolls. Atoll
Research Bulletin 38. Washington: Pacific Science Board.
Schneider, E. (1891) Tagebuchblatter von Jaluit. Deutsche Kolonialzeitung 4, 30-34;46-
48;58-61;75-77.
Spennemann, D.H.R. (1992) Cultural Resource Managment Plan for Majuro Atoll,
Republic of the Marshall Islands. 2 Vols. Washington: U.S.Department of Interior,
Office of Territorial and International Affairs. Part I: Managment Plan 543 pp. Part II:
Appendices 352 pp
Spennemann, D.H.R. (1996) Gifts from the waves. A case of marine transport of obsidian
to Nadikdik Atoll and the occurrence of other drift materials in the Marshall Islands.
Johnstone Centre of Parks, Recreation and Heritage Report N° 23. Albury, NSW.:
Charles Sturt University, The Johnstone Centre of Parks, Recreation and Heritage
Spennemann, D.H.R. and W. R. Ambrose (1997) Floating obsidian and its implications
for the interpretation of Pacific prehistory. Antiquity 71(271): 188-193.
SSL a)" —0
8
Spennemann, D.H.R. and I. G. Marschner (1995) Association between ENSO and
typhoons in the Marshall Islands. Disasters 19(3), 194-197.
Thomas, P.E. (ed.) (1989) Report of the Northern Marshall Islands Natural Diversity and
Protected Areas Survey, 7-24 September 1988. Noumea: South Pacific Regional
Environmental Programme.
Wells, J.W. (1951) The Coral Reefs of Arno Atoll, Marshall Islands. Atoll Research
Bulletin 9. Washington: Pacific Science Board, National Research Council.
ATOLL RESEARCH BULLETIN
NO. 446
DISTRIBUTION OF RAT SPECIES (RATTUS SPP.) ON THE ATOLLS OF THE
MARSHALL ISLANDS: PAST AND PRESENT DISPERSAL
BY
D.H.R. SPENNEMANN
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
OCTOBER 1997
160 162 164 166 168 170 172 174
Q
Eneen-Kio
Bokak
db
i i
! H i H I
© Oe aa ot
i (ej Taka
Ailinginae Ailuk
i Jemo § ‘Mejit i
eal Like py eer
wajalein oo
wm OthO 4 ae
Ujelang | : Sy |
Ujae a
S Maloela
Erikup ] F
+ Lae |
Lib Nenu Aur 8 ;
= Ra og yf sp ae
can Majuro Nae
oS
Ailinglaplap
I Mile
Namdrik Be als - Narikrik |
Kili : ;
Ebon
nautical miles
Figure 1 Index map of the Marshall Islands showing the atolls mentioned in the text.
~ 20
18
16
14
12
10
DISTRIBUTION OF RAT SPECIES (RATTUS SPP.) ON THE ATOLLS OF
THE MARSHALL ISLANDS: PAST AND PRESENT DISPERSAL
by
Dirk H.R. Spennemannt
INTRODUCTION
The study of dispersal processes of small mammals, and especially of rodents, has a wide
range of applications and until recent years there were few publications discussing the
colonisation of ‘oceanic’ islands by small mammals (cf. Crowell, 1986; Diamond, 1987;
Hanski, 1986; Heany, 1986; Lomolino, 1986).
This essay will be concerned with the distribution of rat species in the Marshall Islands
and its implications on the interpretation of the settlement and human use of the atolls. It will
be argued that in all instances the introduction of rats was caused by people and that
accidental transport, such as rafting on drift wood and the like, is as unlikely as introduction
by means of ship wrecks. Human transport as well as the rats’ own inability to cross great
distances of water makes them bad zoogeographical markers, as already pointed out by
Braestrup (1956), but it is precisely this trait that is of concern here. This paper will argue
that the Polynesian rat (Rattus exulans) was an intentional introduction to the area and that
its distribution throughout the Marshall Islands was a deliberate strategy.
THE MARSHALL ISLANDS
The Marshall Islands (Aeon Kein Ad), comprising 29 atolls and 5 islands, are located in the
northwest equatorial Pacific, about 3790km west of Honolulu, about 2700km north of Fiji
and 1500km east of Ponape. With the exception of the two northwestern atolls, Enewetak
and Ujelang, the Marshall Islands are arranged in two island chains running roughly NNW
to SSE: the western Ralik Chain and the eastern Ratak Chain (figure 1). Not counting the
five islands, Jemo, Jabwat, Kili, Lib and Mejit, the atolls of the Marshall Islands range
from very small, with less than 3.5km2, such as Nadikdik (Knox) Atoll, to very large. With
2173km? lagoonal area, Kwajalein Atoll has the distinction to be the atoll with the World’s
largest lagoon. Distances between neighbouring atolls range from as little as 7nm (as in the
case of Nadikidik and Mile) to over 400nm.
There is a range of rainfall regimes, ranging from almost 4000mm yr’! as measured on
the southern atoll of Jaluit (5°47’N) to 1000mm yr"! as noted for the northern atoll of Wake
(19°28’N). Concomitant with that comes a range of vegetation patterns with drier ecotones
prevailing in the north. The lack of a permanent ground water lens (Ghyben-Herzberg lens)
I The Johnstone Centre, Charles Sturt University, PO Box 789, Albury NSW 2640, Australia.
e-mail: dspennemann @csu.edu.au.
Manuscript received 27 June 1997; revised 4 September 1997
SS QS
z
makes these atolls very marginal for human habitation. It is thus not surprising that these
islands have been recorded as uninhabited in the past (cf. Spennemann, 1992). The
question arises whether these islands were ever visited by the Marshallese, either on a
temporary of a semi-permanent basis.
THE RAT SPECIES
Today, there are three rat species present in the Marshall Islands (table 1): the Polynesian rat
(Rattus exulans), the European rat (Rattus rattus, ‘black rat’), and the Norway rat (Rattus
norvegicus). In addition, the house mouse (Mus musculus) is reputedly present on Majuro,
Enewetak and possibly Kwajalein Atolls (Berry & Jackson, 1979). The pre-World War II
rat population of the Marshall Islands, it seems, did not comprise Rattus rattus or R.
norvegicus. Before we review the historic evidence, let us look at the dispersal mechanisms
used by rats.
The Polynesian rat is a fairly sedentary animal with a limited home-range. Contrary to
black rats (Rattus rattus), the Polynesian rat was not observed marking its territories
(Tomich, 1970). It has a predominantly herbivorous diet (Bettesworth, 1972; Fall et al.,
1971) but has also been observed predating on insects (Harrison, 1954; Fall et al, 1971),
snails (Harrison, 1961), land crabs (Moseby et al., 1973), lizards (Crook, 1973; Whitaker,
1973), turtle hatchlings (Balazas, 1983; Hoeck, 1984, p. 242), and bird eggs (Atkinson,
1978; Bourne, 1981; Norman, 1975).
While Polynesian rats can be a plague on European-style monoculture plantations
(Bianchi & Smythe, 1965; Bonin, 1982, 1986; Canter Visscher, 1957; Friend, 1971;
Halafihi, 1985; Pierce, 197la, 1971b; Twibell, 1973; Williams, 1974, 1975, 1982;
Williams & Misikini, 1972; Wodzicki, 1969), they were little problem in the horticultural
framework of the Marshallese culture. Indeed, Chamisso (1986) mentions that the number
of rats had already increased in the period between his first (1816) and second visit (1817)
to Wotje Atoll, destroying most of the plants planted in a model garden. Thus it was decided
to leave behind a number of cats.
It has been put forward that Rattus exulans is responsible for the decline of the lizard
fauna in New Zealand and beyond (Crook, 1973; Morrison, 1954, p. 4; Whitaker, 1973,
1978). Elsewhere it had been argued (Spennemann, 1989, p. 142) that this fact might
explain the observed extinction of large lizard species after initial human settlement of
oceanic islands (Poulsen, 1987; Pregill & Dye, 1989).
Traditionally, that is before the arrival of the first European visitors, the ‘bird atolls’ of
Jemo, Taka, Bikar and Bokak had been regarded as refuges where the taking of birds and
eggs has been tightly regulated by custom (Erdland, 1914; Fosberg, 1957; Tobin, 1952).
The fact that the bird populations continue to thrive (Amerson, 1969; Thomas, 1989) may
indicate that the presence of Rattus exulans is not detrimental to the overall bird population.
Both R.rattus and R.norvegicus are omnivorous and take whatever food is available. In
addition, both species are on the whole substantially more carnivorous than R. exulans and
have been shown to prey not only on insects, but also on bird eggs, bird fledglings, lizards,
3
land snails, molluscs, turtle hatchlings, and land crabs (Atkinson, 1978; Austin, 1948;
Bailey & Sorensen, 1962, Bettesworth & Anderson, 1972; Crook, 1973; Daniel, 1973; Fall
et al., 1971; Harrison, 1961; Ramsay, 1978; Swink ef al., 1970; Watts & Aslin, 1982;
Whitaker, 1973, 1978). The two larger species are also known to displace R.exulans from
their environmental niche (Atkinson, 1973). Once established the rats have been shown to
be quite resilient against natural disasters, being able to survive at least short-time flooding
of an island by storm surges (as evidenced by the tidal surge generated by the Enewetak
nuclear tests; Jackson, 1969).
A local example of the impact of introduced rats comes from Wake Atoll (Eneen-Kio),
where the most dominant mammal on the atoll was the Polynesian rat. During the Japanese
period of occupation in World War II the R. rattus was introduced (Bryan, 1959). The
original bird life consisted of about a dozen different species of sedentary sea birds, and a
few species of migratory sea birds. The only nonmigratory land bird native to and only
occurring on Wake was the flightless rail, Rallus wakensis, which was still seen by the
Tanager expedition in 1922, but which is now presumed to be extinct (Bryan, 1959). Given
the introduction of shipborne rats in Japanese times an eradication of the rail by predatory
rats possibly couped with human predation appears to be the most likely explanation of its
extinction.
DISPERSAL OF RATS
The dispersal of smal] mammals over greater and smaller expanses of water is thought to
have happened in three basic ways:
i) by accidental rafting on material floating in the water (such as tree trunks, logs,
islands of vegetation, and other debris);
11) by swimming; and finally
111) by human-induced transport on boats, ships and rafts.
In the cold climates of the high latitudes movement over frozen lakes and the like, as well as
rafting on ice floats, is also possible (cf. Lomolino, 1986). Accidental rafting on debris
depends entirely on the direction of wind and surface currents and can thus be assessed by
the means of computer simulation studies (cf. Ward et al. 1973) as well as a review of
documented drift voyages. In the Marshall Islands drift has been documented for the
following places of origin: California, North America; Central Solomons; Japan; various
atolls in central Kiribati; Krakatau, Indonesia; Maui, Hawaii; Palmyra Atoll, Line Islands,
Kiribati; Philippines (?); Tuluman I., Bismarck Archipelago, PNG; and Lamotrek,
Pingelap, Woleai and Yap in the Carolines, Federated States of Micronesia (Spennemann,
1996; in press). Internal drift has been documented for the Ratak atolls Mile and Mejit, in
both cases reaching atolls in the Ralik Chain.
Dispersal of small terrestrial animals over long stretches of open water is impeded and
rats have been shown to be unable to cover distances in excess of 2km on their own account
(Jackson & Strecker, 1962; Spennemann & Rapp, 1987, 1989). Survival on drifting items
over prolonged periods of time is also unlikely due to prolonged exposure to the tropical
sun coupled with a lack of water and food. Another argument against successful large-scale
4
accidental dispersal of Polynesian rats on drift wood is the lack of this species on Johnston
Atoll (Amerson & Skelton, 1976) and the French Frigate Shoals (Amerson, 1971), places
which are not known to have had pre-European settlement at any time.
Thus, rats occurring on an isolated atoll are very likely to have been introduced at one
point in time by people either intentionally or accidentally as stowaways. This has also been
assumed previously by some authors (cf. Tate, 1935; Luomala, 1975). Parr (1941, p. 95)
comments that the Polynesian rats on Wake Island were likely to come from wrecks or from
“Polynesian” canoes. Unlike R.rattus and R.norvegicus, which are both known to be
shipboard rats and thus could be accidental European or Asiatic import during the last two
centuries, Polynesian rats are not known to infest vessels.
RATS AS A FOOD SOURCE
An unintentional human introduction of rats to the Marshall Islands is very unlikely, given
the size of Marshallese voyaging canoes which were commonly about 18 to 20m long
(exceptionally up to 30m) and had rather narrow hulls (Alessio, 1990; Browning, 1972;
Chamisso, 1910, 1986; Erdland, 1914; Finsch, 1887; Hambruch, 1912; Hernsheim, 1887;
Kramer, 1905; Kramer & Nevermann, 1938). Given that size, then, rats would have been
noticed if present. Rather, it would appear, rats were a welcomed source of food which —
once released — could fend for itself and thus were taken along as deliberate introductions.
The Polynesian rat is believed to have originated in the Malayan region (Tate, 1935;
Musser & Newcomb, 1983; Roberts, 1991), to have been spread by native canoes, and to
have been deliberately introduced to many islands by Polynesians who considered it a
valuable food source. There is archaeological evidence for pre-European distribution of
Rattus exulans on other atolls and islands in the Pacific, such as Nukuoro (Davidson,
1971); Kapingamarangi (Leach & Ward, 1981); Tikopia (Kirch & Yen, 1982, p. 277);
Kiribati (Luomala, 1975), 'Eua, Tonga (Spennemann, 1987); Ha'apai, Tonga (Dye, 1987;
Spennemann, 1988); Tongatapu, Tonga (Poulsen, 1987); Niuatoputapu, Tonga (Kirch,
1988); and Easter Island (G.Clark pers.comm.).
In Tonga, Polynesian rats have been part of the diet (Gifford, 1929, p. 339; Martin,
1817, p. 279) and hunted for food and for entertainment (Martin, 1817, p. 279-283; Vason,
1810, p. 102-103). Polynesian rats have been seen on numerous now uninhabited islands,
which have later on proven to have carried human occupation, eg. Henderson I. (Schubel &
Steadman, 1989; Sinoto, 1983; Tate, 1935). In the Marshall Islands rats were eaten mainly
by women. Chamisso, for example, observed rats being eaten on Wotje and Uterik in
1816/1817 (Chamisso, 1910, p. 169). Eating rats was common among several Pacific
cultures where pigs (if present) and chicken were reserved for feasts and where terrestrial
animal protein was rare. Eating rats, the only ubiquitous animals around, appears to have
been a convenient means to provide protein for pregnant and lactating women.
DISTRIBUTION OF RATS IN THE MARSHALL ISLANDS
Overall the historic documentation of the rats is limited as they were never the focus of
detailed study until after World War II.
Rattus exulans was observed on Wake (Eneen-Kio) possibly as early as 1568 is the
identification of Wake or Bokak with Alvarez de Mendafia's San Francesco Island is correct
(cf. Hezel, 1983, p. 29; Werstein, 1964, p. 13). Rattus exulans was also observed by the
Russian Exploring Expedition of 1816/17 on Maloelap (Kaven), Wotje and Uterik
(Chamisso, 1910, p. 169, 1986; p. 156). Chamisso comments that some informants
claimed that the rat was nonexistent on Bikar Atoll. This should be read cum grano salis as
Chamisso's informants’ knowledge on the peripheral atolls was very limited at best, not
very surprising in view of that fact that he was not a Marshallese but came from an atoll in
the Western Carolines. It is of significance, however, that in Chamisso's opinion his
informant Kadu could only think of the rats as a companion to people (Chamisso, 1910, p.
169).
Chamisso (1986, p. 156, 196) mentions that the number of rats had already increased in
the period between his first (1816) and second visit (1817) to Wotje Atoll, destroying most
of the plants he had planted in a model garden. Cats were released to act as vermin control,
but by 1830, when Kotzebue returned to Wotje, the number had not diminished (Kotzebue,
L830 ssp. 308):
The U.S.Exploring Expedition saw Polynesian rats in 1840 on then uninhabited Wake
Island and collected some specimens (Cassin, 1858; Peale, 1848; Pickering, 1879; Poole &
Schantz, 1942). The Tanager expedition in 1922 recorded only Rattus exulans for Wake
(Picking, 1922), where they appear to have occurred in reasonable numbers. Following the
establishment of the Pan American Airways station on Wake and the creation of open
rubbish tips, Polynesian rats were to become a plague of major proportions and eventually
were the focus of several eradication campaigns (Anonymous, 1941; Bryan, 1959;
Devereux, 1947; Foulton, 1939; Grooch, 1936; Miller, 1936).
In the late 1880s, with the beginning of the German colonial administration, the number
of scientific studies increased, mainly focussing on the avifauna, as rodents were seen as a
pest (Anonymous, 1895) and not the focus of enquiry. As a side-effect of increased copra
production the number of rats increased too. The German district Officer Georg Merz,
stopping at Majuro Atoll in 1910 on occasion of his annual inspection voyage, reports on
large numbers of rats in plague proportions and suggest the release of cats to reduce the rat
problem (Merz 1910). The data in hand suggest that the pre-World War II rat population of
the Marshall Islands comprised neither Rattus rattus or R. norvegicus, with the possible
exception of Jaluit and Majuro Atolls, the former the administrative centre of the German
and (later) Japanese Colonial Administrations, and the latter an atoll with a well established
trading station replete with pier.
INTRODUCTION OF RATS, 1885 TO PRESENT
Inter-atoll communication in the Marshall Islands was previously upheld solely by the
means of local canoe transport. Local communication between the atolls, however, seems to
have been largely restricted to the southern part of both the Ralik and the Ratak chains, and
between the southern parts and northern parts of either chain. An investigation of the
distribution of introduced epidemics clearly documents this pattern. For example, Steinbach
(1893), discussing the spread of a syphilis epidemic, mentions that it was prevalent in
Majuro, Ebon and Jaluit Atolls but occured only in limited proportions in the northern
atolls, which had little communication with the former.
With the increasing presence of European traders, however, European vessels and even
ship-/boat-building of European-type vessels, built by J.de Brum on Likiep Atoll, became
more common. Conversely, the inter-atoll transport was increasingly conducted with larger,
European-type vessels (cf. Linckens, 1912). During the period of the German colony, the
Jaluit Gesellschaft operated a steam vessel as well as a number of sailing schooners in the
islands. Further transport was provided by a vessel of the Australian Trading Company
Burns Philp and Co. Apart from the inter-atoll trade, the Jaluit Gesellschaft also operated
“long-distance” voyages to Pohnpei, Palau and New Guinea, In addition, there were the
regular annual visits of German naval vessels. With the exception of Jaluit and Majuro none
of the atolls had proper landing bridges or piers during the German period, and thus all
vessels had to anchor in the lagoon with all trade being conducted by launch or canoe. The
same applies to the few whalers that came in the 1880s to replenish their stores of water and
food (Langdon, 1978, 1979).
Such conditions, however, are not at all conducive to the introduction of shipboard rats.
The same pattern continued during the Japanese period until in the late, 1930 piers were
built on islands earmarked for future military development (Yanaihara, 1940; Japanese
Government, 1929).
The German government introduced quantities of soil to Jaluit to run the experimental
garden. The import occurred mainly in the form of ship's ballast, brought by copra trading
vessels returning partially empty from the volcanic high islands in the Carolines (such as
Ponape) (cf. Anonymous, 1895; Fosberg, 1961; Fosberg & Sachet, 1962, p. 1; Stevenson,
1914, p. 150). It is possible that rats were also ‘landed’ during the unloading of these
vessels.
The Japanese have a history of both unintentional and intentional introductions: during
the period of Japanese administration import of soil directly from Japan has been reported
(Price, 1935, p. 256). The Japanese, intent on staying for a long time, imported night soil
from Japan to improve the soil on both Wake and Wotje Atolls (Kephardt, 1950, p. 34).
Import of the same material can be assumed for two, or three other major Japanese bases,
namely Kwajalein/Roi-Namur, Taroa (Maloelap Atoll) and Jaluit, all of which had been
duilt before the begin of the Pacific War. These soil imports are likely to have been very
small, just confined to gardening plots.
Table 1. The occurence of Varanus indicus and the distribution of rodent species on the
atolls of the Marshall Islands. [1]
Rattus | Rattus | Rattus Mus Japanese | Varanus
Atoll exulans rattus norvegicus | musculus | Development indicus
Ailinginae a lt
Ailinglaplap
| Ailuk
Arno
Aur
Bikar
Bikini
Ebon
Eneen-Kio (|
Enewetak
Erikup
Jabwat
Bile B(3) | a major Present
Kwajalein
Lae
a major Eradicated
a a a i major Eradicated
Majuro
Maloelap
Mejit
Milli
Nadikdik
Namorik
| Namu
Rongelap
Rongerik
Taka
Taongi
| Ujae
Ujelang
BREE ~ BE ~~ ee ~ 2 e~
a
@
4
2.
le)
=
hele
|} major Eradicated (?)
[1] Compiled after Berry & Jackson 1970; Betlack & Eckhardt 1945; Bryan 1959; Cassin 1858; Chamisso
1986; Finsch 1893; Fosberg 1955, 1956; 1957, 1990; Gressitt 1961; Hatheway 1953; Kotzebue 1830;
Marshall 1950; 1957; Thomas 1989; and own observations. [2] Wake Atoll in US parlance. [3] Introduced
by the U.S. forces after 1944. [4] Introduced by the Japanese in the 1930s.
Apart from introducing plant pests along with the soil, the Brahminy blind snake
(Ramhotyphlos brahmina, TYPHLOPIDAE) seems to have been introduced, occurring so far
only on Enewetak Atoll (but there on different islands). The secretive, nocturnal and earth
burrowing nature of this harmless snake makes its discovery a difficult (Lamberson, 1987).
During the Japanese occupation of Wake in World War II (Dec.1941—Sept.1945), Rattus
rattus was introduced with devastating effects on the birdlife (Fosberg, 1959). Cunningham
(1961, p. 87), Commanding Officer of the Wake I. garrison and commenting on the events
of December 1941, mentions “Wake Island’s stunted rats’, which seems to refer to the
Polynesian rat, suggesting that the black rat and the Norway rat had not yet arrived.
As these rats were present after the war, their import must have occurred during
Japanese times. Already in the 1930s the Japanese had introduced the brown rat to Wotje
(Marshall, 1950, p. 23) and Jaluit. While Marshall suggests that R. rattus may have been
also introduced to Arno before the war, it is more likely that the landing boat activity of
U.S. forces during the relocation of Marshallese from various Japanese-held atolls via
Majuro to Tutu Island on Arno Atoll (Richard, 1957), is responsible for its introduction.
The distribution of rat species in the Marshalls (table 1) shows that R. rattus and
R.norvegicus are present on those atolls that were major Japanese military installations
during World War II.
Figure 1. A specimen of Varanus indicus caught on Majuro Atoll, Marshall Islands during
1944/1945 (after Betlack & Eckhardt 1945).
9
The rat problem on some bases reached such proportions that Varanus indicus were
introduced to prey upon the rats. Instead, according to local Marshallese informants, the
reptiles predated on the chickens as well as other birdlife. Varanus indicus has been
described for Enewetak (Lamberson, 1987), where an extensive natural history assessment
has been carried out. Immediately after the Pacific War it was found on Majuro when the
US forces occupied the atoll (Betlack & Eckhardt, 1945). Today, Varanus indicus 1S
occasionally caught on Enewetak and brought to the population centres of Majuro and
Enewetak as a pet (pers. obs.).
Figure 2. A specimen of Varanus indicus caught on Enewetak Atoll and brought as a pet to
Majuro Atoll, Marshall Islands (November 1992).
The distribution of the rat species in 1991/92, as shown in table 1, is based on a literature
survey, as well as my own observations. The lack of R.rattus/R.norvegicus on most atolls
is confirmed by own and other observations. Even though no trapping was carried out
where R.rattus/R.norvegicus were present, such as on Taroa (Maloelap Atoll), Mile (Mile
Atoll) and Wotje (Wotje Atoll) they were common and could be observed scurrying
fearlessly on the ground. The rats permitted quite close observation before they ran way.
This is also confirmed by members of the Independent Nationwide Radiological Study that
took radioactivity measurements on all atolls of the Marshall Islands (Simon pers. comm).
The current distribution of R.rattus/R.norvegicus is not an artifact of selective or differential
observation and reporting.
10
POTENTIAL IMPACT OF SHIPBORNE RATS
Given the overall urban and agricultural/horticultural development of the atolls of the
Marshall Islands the few bird atolls remain ecological refuges and sea-bird nesting colonies
of Pacific-wide significance. Any landing of shipborne rats on board of a stricken vessel is
likely to constitute an ecological catastrophe. And shipwrecks, especially of Japanese
fishing vessels, are not uncommon (Spennemann, 1991; Thomas, 1989).
However, not all shipwrecks on atolls necessarily introduce rat species. It is possible to
compile from the literature quite an extensive list of shipwrecks which occurred in the
Marshall Islands over the past 100 years. Yet, none of these vessels introduced any Rattus
rattus and Rattus norvegicus; given the nature of some of the vessels it is highly unlikely
that at least some would not have had rats on board (cf. Hezel, 1979). It would appear that
the wrecks had all been stranded at locations where the rats could not get ashore or where
they died in the surf when the vessels broke up.
The only clear evidence of colonisation by Rattus rattus and Rattus norvegicus in the
Marshalls occurred when ships were moored at piers and where the rats had the chance to
run down mooring lines or gangways. The dispersal of R.rattus/R.norvegicus 1s poised to
increase as piers to unload the field-trip ships or fishing bases have now been constructed
on many atolls. To contain the spread of these two species care needs to be exercised with
lines being properly fitted with regulatory rat disks.
In order to avoid the accidental landing of shipborne rats on the bird atolls, however,
extreme precautions need to be taken, both in view of landing or beaching any support
vessels and in view of the unloading and lightening of the stricken (fishing) vessel.
BIBLIOGRAPHY
Alessio, D. F. 1990. The Likiep documentary. The Likiep taburbur. Waan Aelon Kein
Project Report N° 3. Majuro, Marshall Islands: Alele Museum.
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ATOLL RESEARCH BULLETIN
NO. 447
A POSSIBLE LINK BETWEEN CORAL DISEASES AND A CORALLIVOROUS
SNAIL (DRUPELLA CORNUS) OUTBREAK IN THE
RED SEA
BY
ARNFRIED ANTONIUS AND BERNHARD RIEGL
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
OCTOBER 1997
A POSSIBLE LINK BETWEEN CORAL DISEASES AND A
CORALLIVOROUS SNAIL (Drupella cornus) OUTBREAK IN THE RED SEA
by
Arnfried Antonius! and Bernhard Rieg!!
ABSTRACT
In April-May and in September 1996, a total of 25 reefs were studied between
Taba and Ras Mohammed in the Gulf of Aqaba, Red Sea. In only four of these reefs
Drupella cornus showed up in the transects in low numbers and coral diseases were
found at a moderate level on most reefs. Only the reefs of Ras umm Sidd, near Sharm el
Sheikh, exhibited Drupella cornus as well as coral diseases both at abundant or even
epidemic levels. There definitely seems to be a correlation between abundance of snail
and diseases, but the question of "what comes first" remains to be investigated : does
massive coral die-off (mostly White Syndromes) attract or benefit Drupella cornus and
thus promote a population explosion, or does a massive D. cornus invasion promote an
epidemic of White Syndromes on corals ?
INTRODUCTION
In the course of a large scale ecological investigation of Gulf of Aqaba coral reefs,
with special emphasis on coral health, the distribution of coral diseases and the impact of
predators was investigated. We detected an abnormally high proportion of dead corals at
Ras umm Sidd (fig.1) and found two major causes contributing to coral mortality:
1) an abundance of coral diseases, mainly White Syndromes (Antonius 1995a), and
2) a population explosion of the corallivorous gastropod Drupella cornus (plate).
The coral species most frequently affected was the branching species Acropora
hemprichi, an important and dominant species on Red Sea reefs (Rieg] & Velimirov
1994). Of all the reef sites studied throughout the Sinai-side of the Gulf of Aqaba (and in
the past also Haq]: Antonius 1988), Ras umm Sidd was found to be the only site
showing this combination of high levels of Drupella-predation associated with high levels
of coral diseases. This observation led us to the question whether the two phenomena
were correlated. Intensive Drupella predation (without associated diseases), that caused
the destruction of wide reef areas, has been reported by others from the northern and
central Red Sea (Schuhmacher 1992; Schuhmacher et al. 1995), from Japan and the
Philippines (Moyer et al. 1982), as well as from Western Australia (Turner 1992, 1994).
1 Institut fiir Palaontologie der Universitat Wien, Geozentrum, Althanstrasse 14,
A-1090 Wien, Austria.
Manuscript received 8 August 1997; revised 2 September 1997
Coral diseases (without associated D. cornus) degrading reef health have been
reported from Caribbean (Antonius 1977, 1981), as well as Indo-Pacific locations
(Antonius 1984, 1988).
The first stage of this study was carried out by both authors in April-May 1996;
later work was conducted by Antonius in September of the same year.
Sharm el
Moija
Pharaoni Beach
- a
The Temple windward
leeward
Figure 1 —_‘ The four survey sites of the study: the fringing reefs of 1) Ras umm Sidd
windward side, 2) Ras umm Sidd leeward side, 3) Pharaoni Beach, and the patch reefs
of 4) The Temple.
MATERIALS and METHODS
Coral diseases and other coral destroying agents encountered during this survey
were the following: Black Band Disease (BBD), Black Overgrowing Cyanophyta
(BOC), White Band Disease (WBD, Tissue Bleaching (TBL), and Shut-Down-Reaction
(SDR); all listed and described in Antonius (1995a), as well as a newly discovered
Skeleton Eroding Band (SEB) which is presently under investigation. WBD, TBL, and
SDR are jointly referred to as White Syndromes (WS). Also recorded was the coral-
eating snail Drupella cornus (DRU).
The semi-quantitative Belt Method (Antonius 1995b) was used to assess these
syndromes. It is a time-count technique using one half-hour of observation time, which is
considered aSCAN. During every scan, the diver swims fairly close to the reef surface
and notes down all pathologic syndromes on corals that are encountered. The numbers of
syndromes counted during one scan are arranged in categories: 1-3 cases = condition 1,
rare, 4-12 cases = condition 2, moderate; 13-25 cases = condition 3, frequent; 26-50
cases = condition 4, abundant; 51-100 cases = condition 5, epidemic, and any number
in excess of 100 = condition 6, catastrophic. Four sites were surveyed this way: Ras
umm Sidd windward, Ras umm Sidd leeward, Pharaoni Beach, and The Temple (figure 1
and table 1).
A similar semi-quantitative method was used to assess the impact of Drupella
cornus predation on the local populations of A. hemprichi.. We sampled the same sites
except Pharaoni Beach (fig. 2). During a 30 minute dive, which followed a depth gradient
to 25 m depth, D. cornus populations were assessed in the same way described above
(Belt-Method). In addition to that, all A. hemprichi colonies were recorded and grouped
into four categories:
- alive, meaning no signs of recent partial mortality (no white areas)
- dead, meaning recently dead (whole colony white)
- partly dead, with some recently denuded branches (white skeleton)
- discolored, some colonies did not display the typical blue or green color, but were of a
faded yellow and had filamentous algae settling on the tissues, which indicated poor
health. This category was used in order to check for sources of mortality other than
Drupella.
RESULTS
Of 25 reefs studied between Taba and Ras Mohammed in April-May 1996
(Antonius 1996, Rieg] 1996), Drupella cornus was found on most of the surveyed reefs,
but in densities so low that they did not always show up in a scan (few branches on
digitate colonies stripped of tissues, no freshly dead entire colonies). At the level of
condition | (rare), Drupella cornus was found on four reefs (Marsa el Muqabila, Nabq,
Turtle reef, Kashaba beach: some stripped branches, also few completely stripped
colonies, but less than 10% of digitate colonies affected).
100
A. hemprichi at Ras umm Sidd, windward
% of
all
colonies
alive dead partly dead discoloured
100
A. hemprichi at Ras umm Sidd, leeward
% of
all
colonies
alive dead partly dead discoloured
100
A. hemprichi at "The Temple"
tS
% of
all 50
colonies
25
NNNNASNSNSNSANASN
‘N
NANANSNSSNANSNS
NNANNNSSNSNASNS
NANNANNSASNASNS NNANNNSNSASNSSNS
DS
alive dead partly dead discoloured
Figure 2 The state of health of Acropora hemprichi populations at three
sample sites of the study area.
Plate Underwater photograph showing Drupella cornus feeding on a branch of
Acropora hemprichi at the exact borderline of an active White Band Disease.
In very high densities, up to condition 5, D. cornus was only found at Ras umm
Sidd (frequent completely stripped colonies, most colonies with stripped branches, over
20%, and up to 60%, of all colonies affected). These are elevated levels even when
compared to values of high Drupella frequency obtained by Schuhmacher et al (1995)
from Aqaba (1% coral death on average for the whole shallow reef, up to 30% tissue
depletion).
The reefs with low and medium Drupella abundance also had a normal incidence
of coral diseases as diagnosed according to Antonius (1995a). Ras umm Sidd and the
adjacent Pharaoni Beach toward Ras Katy (fig. 1), however, showed high levels of coral
diseases. They increased in frequency from the windward side of Ras umm Sidd to its
leeward side and increased even further towards Pharaoni Beach and The Temple. Coral
diseases were censused on the reef flat, on the reef edge and slope, and on the patch reefs
immediately in front of the fringing reef (The Temple). The general health status of the
reef therefore decreased markedly towards the center of the bay (table 1).
Table 1
Occurrence of coral diseases and Drupella cornus damage, as well as their frequency
(= condition-numbers) at the four sample sites :
WBD (White Band Disease) )
TBL_ (Tissue Bleaching) ) = WS (White Syndromes)
SDR_ (Shut-Down-Reaction) )
BOC (Black Overgrowing Cyanophyta),
SEB (Skeleton Eroding Band),
DRU (Drupella cornus)
Condition he eraike (1-3 cases per scan)
2 = moderate (4-12 cases per scan)
3) = irequent (13-25 cases per scan)
4 = abundant (26-50 cases per scan)
Se epidennic (51-100 cases per scan)
6 = catastrophic (above 100 cases per scan)
Ras umm Sidd
Windward BOC WBD TBL
flat + slope 3 Di ee
Ras umm Sidd
Leeward
BOC’ WED TBO SDR” DRU SEB
flat 4 4 3 2 1
slope 4 - 3 2 5 1
Pharaoni Beach BOC WED TBE ‘SDRe"*DRU SEB
5 4 4 4 4 2
The Temple BOC "WBD° TBE’ SDR’ “DRU“SEB
5 5 + 4 SS) 2
Similar to the distribution of coral diseases, the frequency of Drupella cornus also
increased from the windward side of Ras umm Sidd toward the bay (table 1). At the
windward side of Ras umm Sidd, the shallow (0.5 m) reef-flat as well as the (steep) fore-
reef-slope were practically free of Drupella (DRU). On the leeward side, the frequency of
pathologic syndromes increased, but only on the slope was it accompanied by Drupella
(table 1). At both sites, Pharaoni Beach and The Temple, incidences of coral diseases and
Drupella frequency increased even further (table 1), with Drupella occurring below 1-
1.5 m at Pharaoni Beach and below 3m at The Temple.
Although observation time was too short to obtain absolute certainty, we were able
to distinguish three phases of WS-DRU interaction:
Phase 1: Drupella cornus snails, when occurring in low numbers, are usually
feeding on the exact interface of a WBD (fig. 3); with such an open wound available, they
do not attack healthy coral tissue.
Phase 2: Larger concentrations of D. cornus are feeding on healthy coral tissue at
a speed far exceeding that of a WBD; this 1s the situation most frequently encountered at
Ras umm Sidd.
Phase 3: The impact of excessive feeding by Drupella triggers a SDR,
destroying more coral tissue than is occupied by snails; large numbers of D. cornus, now
stranded on a coral branch without tissue, move on to new feeding grounds.
During a re-survey in September 96 it was noted that the categories of conditions,
established in April-May, were changing. For example, when a condition 4 (abundant) in
April-May covered roughly 30 cases, in September the number of cases had increased to
between 40 and 50.
DISCUSSION
This correlation of declining reef health and increasing frequency of Drupella led
us to the question how these two independent phenomena, i.e. WS diseases and D. cornus
predation, could become connected through such a "circulus vitiosus" at Ras umm Sidd ?
Ras umm Sidd is one of the most frequented diving sites in the northern Red Sea
(Hawkins & Roberts, 1992). Two hotels have private beaches inside the bay between Ras
umm Sidd and Ras Katy (one of them the sample site Pharaoni Beach). Furthermore, it is
close to the town of Sharm el] Sheikh and its busy port at Sharm el Moija.
Reasons for the decline in reef health could be related to the leaching of toxic
substances from the antifouling of ship bottoms, the disposal of sewage and septic tanks
from the dive boats, and also to considerable impact by divers and swimmers. Tourists
coming in by boat are usually told by their dive-guides what to avoid under water. But not
everybody heeds the advice, and many divers come from land. The result is that corals
are constantly being touched and stepped upon, thus remaining in a state of chronic
irritation.
Whatever the reasons for the bad health particularly in the center of the bay, only
this already weakened reef showed signs of a Drupella outbreak, while other reefs in the
area showed normal health and low frequency of Drupella. At Ras umm Sidd, the
frequency of coral diseases increased towards the bay both on the reef flat, where no
Drupella were encountered, and the reef slope, where Drupella were frequent (table 1).
Therefore, on the reef flat, there is little room for misidentifying White Syndromes as
Drupella damage (table 1, leeward, flat). On the reef slope this situation was not so clear,
as Drupella damage and White Syndromes are both frequent in this zone (table 1,
leeward, slope) and not always easy to distinguish.
However, there is evidence that Drupella and WS occurrence may be linked in
some way. For example: when a large WBD stretches across a corallum and a scant few
specimens of Drupella are feeding on the exact interface of the disease (= phase 1), they
were apparently attracted by the disintegrating coral tissue. The same phenomenon was
observed involving the "fireworm" Hermodice caruncullata in the Caribbean Sea
(Antonius 1975). A situation comparable to phase 2 (as defined in "Results") has been
observed in the behavior of the crown-of-thorns starfish Acanthaster planci in the past
(Antonius 1971). And a phase 3 phenomenon. i.e. a SDR outrunning the predator, was
originally also documented for Hermodice caruncullata (Antonius 1977).
Thus, this particular sequence of events seems to be reasonably clear. However,
since we do not know whether every local D. cornus invasion originated at the site of a
WBD, the basic question still remains: does massive coral die-off (mostly WS) benefit
Drupella cornus and thus promote a population explosion, or does a massive Drupella
invasion promote epidemic White Syndromes on corals ?
ACKNOWLEDGEMENTS
The beginning stages of this work were partly a result of contract SEM/03/220/025
A - Egypt with the Egyptian Environmental Affairs Agency. Following investigations
were supported by project No. P11734-BIO of the Austrian Science Foundation (FWP).
The collaboration of R.F.G. Ormond and M.P. Pearson is gratefully acknowledged. E.
Sadallah, A. Mabrouk and K. Mahmoud assisted with data gathering.
REFERENCES
Antonius A (1971) Das Acanthaster Problem im Pazifik. Internationale Revue der
gesamten Hydrobiologie, 56 (2): 283-313.
Antonius A (1975) Predation by the Polychaete Hermodice caruncullata on Coral Reefs.
In: Field Guide to some Carbonate Rock Environments. Ed. H.G. Multer,
Kendall Hunt Publishing Comp. Revised Edition, pp. 133K-133M.
Antonius A (1977) Coral Mortality in Reefs : A Problem for Science and Management.
Proc. Third Internat. Coral Reef Symp., University of Miami, Florida, 2: 618-623.
Antonius A (1981) The "Band" Diseases 1n Coral Reefs. Proc. Fourth Internat. Coral
Reef Symposium, University of the Philippines, Manila, 2: 7-14.
Antonius A (1984) Coral Diseases in the Indo-Pacific: A First Record. P.S.Z.N.I: Marine
Ecology, 6(3): 197-218.
Antonius A (1988) Distribution and Dynamics of Coral Diseases in the Eastern Red Sea.
Proc. Sixth Internat. Coral Reef Symp., J.C. Univ., Townsville, Australia, 2: 293-
298.
Antonius A (1995a) Pathologic Syndromes on Corals: a Review. Publ. Serv. Geol. Lux.,
23, Proc. 2nd Europ. Regional Meeting ISRS: 161-169
Antonius A (1995b) Coral Diseases as Indicators of Reef Health: Field Methods. Publ.
Serv. Geol. Lux., 23, Proc. 2nd Europ. Regional Meeting ISRS: 231-235
Antonius A (1996) Sinai Coral Reef Health Survey. Report to the National Park Service
of Egypt, 29 pp.
Hawkins J, Roberts C (1992) Can Egypt's Coral Reefs Support Ambitious Plans for
Diving Tourism? Proc. 7th Int. Coral Reef Symp., Guam 1992, Vol. 2: 1007-1013
Moyer JT, Emerson WK, Ross M (1982) Massive Destruction of Scleractinian Corals by
Muricid Gastropods, Drupella, in Japan and the Philippines. The Nautilus 96:69-82
Rieg] B (1996) Design of a Coral Reef Monitoring System and Establishment of a
Baseline Database for the Ras Mohamed National Park: Pilot Monitoring Project,
(in press).
Riegl B, Velimirov B (1994) The Structure of Coral Communities at Hurghada in the
Northern Red Sea. PSZNI Marine Ecology 15 (3/4): 213-233
Schuhmacher H (1992) Impact of Some Corallivorous Snails on Stony Corals in the Red
Sea. Proc. 7th Int. Coral Reef Symp., Guam, 1992, Vol. 2: 840-846
Schuhmacher H, Kiene WE, Dullo WC (1995) Factors Controlling Holocene Reef
Growth: An Interdisciplinary Approach. Facies 32: 145-188
Turner S (ed.) (1992) Drupella cornus: A Synopsis. CALM Occasional Paper No.3, 104
Pp.
Turner S (1994) Spatial Variability in the Abundance of the Corallivorous Gastropod
Drupella cornus. Coral Reefs 13: 41-48
ATOLL RESEARCH BULLETIN
NO. 448
MARINE ALGAE FROM OCEANIC ATOLLS IN THE SOUTHWESTERN
CARIBBEAN (ALBUQUERQUE CAYS, COURTOWN CAYS, SERRANA BANK,
AND RONCADOR BANK)
BY
GUILLERMO DIAZ-PULIDO AND GERMAN BULA-MEYER
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
OCTOBER 1997
MARINE ALGAE FROM OCEANIC ATOLLS IN THE SOUTHWESTERN
CARIBBEAN (ALBUQUERQUE CAYS, COURTOWN CAYS, SERRANA BANK,
AND RONCADOR BANK)
BY
GUILLERMO DIAZ-PULIDO* and GERMAN BULA-MEYER**
ABSTRACT
A total of 171 taxa of benthic marine algae are recorded from four oceanic atolls
in the southwestern Caribbean Sea (Albuquerque Cays, Courtown Cays, Roncador Bank
and Serrana Bank). The algae were collected in the different geomorphological zones and
bottom habitats occurring in these reef-complexes, and within a depth range from
intertidal to 40 m. Of the total taxa found, 6 are Cyanobacteria, 61 Chlorophyta, 22
Phaeophyta and 82 Rhodophyta. Twenty seven taxa are new records for the Colombian
Caribbean. The marine flora of these atolls is closely related with that of the northern
Caribbean phytogeographical region.
INTRODUCTION
Albuquerque and Courtown Cays and Serrana and Roncador Banks are four small
atolls part of the Colombian Archipelago of San Andrés and Providencia in the
southwestern Caribbean Sea (Fig. la and 1b). Several phycological surveys have been
carried out in these remote reefal areas, including studies on algal distribution (Hay, 1984;
Diaz-Pulido and Diaz, in press) and records of some species from this area (Littler and
Littler, 1992; Bula-Meyer and Diaz-Pulido, 1995). However, to date, no intensive floristic
accounts have been published from these atolls, contrasting with the numerous checklists
of benthic marine algae available from other southwestern Caribbean localities [i. e., San
Andrés and Providencia islands (Kapraun, 1972; Schnetter, 1976, 1978; Marquez, 1992),
Belize (Norrris and Bucher, 1982; Littler et al., 1995), Glovers Atoll (Tsuda and Dawes,
1974), Swan Islands (Taylor, 1975), and Miskito Bank (Phillips et al., 1982)]. The
checklist presented here contains additional information about the different environments
where algae were collected. This list may be useful for future biogeographical studies, and
represents a further step in marine biodiversity studies of this little known Caribbean area.
*Instituto de Investigaciones Marinas y Costeras, INVEMAR, A.A. 1016, Santa Marta,
Colombia, South America.
**Universidad del Magdalena, Departamento de Biologia, A.A. 890, Santa Marta,
Colombia, South America.
Manuscript received 12 October 1996; revised 29 May 1997
STUDY AREA AND METHODS
A full description of the study area and characteristics of the marine habitats, as
well as notes on geologic origin, and climatic and oceanographic conditions can be found
in Milliman (1969), Geister (1992) and Diaz et al. (1995; 1996). All four atolls exhibit in
general the same basic geomorphological features and marine habitats, presenting each a
windward and leeward fore-reef terrace, continuous peripheral reefs (“barrier reef’), a
lagoonal terrace and discontinuous peripheral reefs on the leeward side, resulting in rather
open lagoons (Fig. la and 1b). The algae were collected by the first author during two
cruises aboard the R/V Ancon (INVEMAR, Santa Marta, Colombia) conducted in May-
June 1994 to Courtown (12° 24' N, 81° 28' W) and Albuquerque (12° 10' N, 81° 51' W),
and to Serrana (14° 16' N, 80° 20' W) and Roncador (13° 34' N, 80° 04' W) one year later.
A total of 111 collecting sites were visited (28 at Courtown, 28 at Albuquerque, 29 at
Serrana and 26 at Roncador), comprising the various geomorphological zones and marine
habitats down to a depth of 40 m (Table 1). Collected material were fixed in 4 % formalin
and mounted on herbarium sheets. Wet specimens were later transferred to 70 % alcohol
and deposited in the algal collection of INVEMAR; dry specimens were deposited in the
herbarium of the first author (DP). The taxonomic arrangements proposed by Wynne
(1986) were mainly followed. Some groups were identified with specialized literature:
Cyanobacteria (Humm and Wicks, 1980; Golubic and Focke, 1978; Drouet, 1981),
Udotea (Littler and Littler, 1990), Avrainvillea (Littler and Littler, 1992) and Dictyota
(Hornig et al., 1992). Each taxa is followed by codes indicating the geomorphological
zone, marine habitat and depth range where they were collected in each atoll. Taxa
reported for the first time for Colombia are preceded by asterisks (*).
RESULTS AND DISCUSSION
A total of 171 algal taxa (162 species, 1 subspecies, and 8 forma) were recorded
from the coral reef complexes of Albuquerque (111 taxa), Courtown (107 taxa), Roncador
(88 taxa) and Serrana (98 taxa); they included 6 Cyanobacteria, 61 Chlorophyta, 22
Phaeophyta and 82 Rhodophyta (Table 2). The families with the highest species numbers
were Udoteaceae (25), Corallinaceae (23), Ceramiaceae (22), and Dictyotaceae (15),
whereas species-rich genera were Dictyota (10), Halimeda (8), Caulerpa (7), and
Avrainvillea, Udotea and Peyssonnelia (6 species each). An analysis by functional-form
groups of macroalgae (Littler et al., 1983 a, b) showed that groups with less complex
morphologies, such as the Filamentous-, Coarsely Branched- and Sheet-Groups, exhibited
the highest number of taxa (39, 37 and 29 respectively), comprising the 64 % of the total
recorded (excluding Cyanobacteria). On the other hand, functional-form groups
comprising tougher macrophytes yielded lower numbers of taxa (e.g. Thick Leathery with
19, Jointed Calcareous with 17 and Crustose with 24). However, a great number of taxa of
calcareous algae (which have examples in different functional-form groups) was recorded
from these reefal areas (59), supporting the argument that they represent a highly diverse
group in environments exposed to great grazing pressures (Littler and Littler, 1984).
3
The most common species was Dictyota cervicornis, occurring in almost all
explored habitats. Other fairly common species were Halimeda opuntia, Lobophora
variegata, D. pfaffii and Amphiroa fragilissima, all of them widely distributed in almost
every environment. Certain algae occurred almost exclusively in specific habitats. Such is
the case of Hydroclathrus clathratus, Acanthophora spicifera, Padina jamaicensis,
Avrainvillea rawsonii, A. digitata, Enteromorpha lingulata, Chaetomorpha gracilis,
Cladophora dalmatica, which were preferentially encountered on shallow flat bottoms of
the lagoonal terrace. Likewise, Rhipocephalus phoenix, Halimeda incrassata, H.
simulans, H. monile and most species of Udotea, Avrainvillea and Penicillus are mainly
dwellers of sand-plain bottoms of the lagoonal basin.
Twenty-seven taxa are new records for the Colombian Caribbean flora:
Oscillatoria lutea C. Agardh, O. submembranacea Ardissone et Strafforella, Phormidium
hendersonii Howe, Schizothrix mexicana Gomont, Calothrix crustacea Schousboe et
Thuret, Struvea ramosa Dickie, Derbesia cf. marina (Lyngbye) Kjellman, D.
vaucheriaeformis (Harvey) J. Agardh, Avrainvillea asarifolia f. olivaceae Littler et
Littler, A. digitata Littler et Littler, A. /evis f. translucens Littler et Littler, A. silvana
Littler et Littler, Udotea cyathiformis v. cyathiformis f. infundibulum (J. Agardh) Littler et
Littler, U. dixonii Littler et Littler, U. Jooensis Littler et Littler, U. Juna Littler and Littler,
Neomeris mucosa Howe, Sargassum histrix J. Agardh, Liagora norrisiae Abbott,
Titanoderma bermudense (Foslie et Howe) Woelkerling, Chamberlain et Silva, T.
prototypum (Foslie) Woelkerling, Chamberlain ef Silva, Botryocladia pyriformis
(Bergesen) Kylin, Balliella pseudocorticata (Dawson) D. Young, Ceramium rubrum
(Hudson) C. Agardh, Griffithsia heteromorpha Kitzing, Lejolisia cf. mediterranea
Bornet, and Hypoglossum caloglosoides Wynne et Kraft. Although these are widely
distributed species in the Caribbean, their distribution in the atolls is restricted to a few
habitats (Table 2).
Habitats in which greater number of stations were established, yielded the higher
number of species [i.e. lagoonal patch reefs of Montastraea spp. (97 species), windward
fore-reef terraces (92 species), and Leeward terraces (81 species)]. Therefore, it does not
appear relevant to make comparisons of species richness between habitats differing in
collecting intensity. Although similar number of stations were visited in the four atolls, a
lesser number of taxa were recorded in Roncador and Serrana than in Albuquerque and
Courtown. This difference may reflect a more intensive collecting effort in the latter
atolls, rather than differences in species richness among them. However, in general terms
it may be stated that the algal flora is quite similar in the four atolls.
The marine algal flora of the atolls studied is relatively rich in species when
compared with other well studied Caribbean reef-complexes: Belize (284 taxa, Norris and
Bucher, 1982; Littler et al., 1995), Swan Islands, Honduras (51 species, Taylor, 1975),
Glovers Atoll, Belize (100 species, Tsuda and Dawes, 1974), Miskito Bank, Nicaragua
(99 species, Phillips et al., 1982), San Andrés Island, Colombia (96 species, Kapraun,
1972; Schnetter, 1976; 1978; 1980), Providencia Island, Colombia (50 species, Marquez,
4
1992), Rosario Islands, Colombia (145 species, Bula-Meyer et al., 1993), Curacao (142
species, Van den Hoeck et al., 1975). Our results agree well with Taylor (1975), that the
phytogeographic relationship of the archipelago of San Andrés and Providencia
(including the atolls studied) to the flora of the northern Caribbean is rather clear.
The list presented here should be regarded as preliminary, since floristic accounts
and collecting were more intensively conducted on hard substrata and areas with luxuriant
coral reef development than in back reef flats and sand and rubble plains, which take up a
significant area in all four atolls. Further collecting is needed to more comprehensively
inventory the marine algal flora of southwestern Caribbean Atolls.
ACKNOWLEDGEMENTS
The first author is indebted to J.M. Diaz, J. Garzon-Ferreira, L.S. Mejia, J.A.
Sanchez, S. Zea, and the crew of the R/V ANCON for their help during the field work.
We thank J.M. Diaz for his review of the english version of the manuscript. Constructive
comments by the reviewers improved this contribution and are much appreciated. Thanks
also go to Dr. R. Schnetter for the revision of some specimens of Dictyota spp. This paper
contains portions of a B.Sc. thesis submitted by GD-P to obtain the Marine Biology
degree at Universidad Jorge Tadeo Lozano, Bogota and Santa Marta, Colombia. This
study was supported by INVEMAR and COLCIENCIAS (Grant No. 2105-09-023-93).
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6
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Table
Codes employed to
geomorphological zones of each algal taxa in Table 2.
CODE
DEPTH
RANGE (m)
8-12
12.1-16
16.1-20.5
1-3.5
6-10
10.1-14
14.1-18.5
30-35
35.1-40
MARINE HABITAT
Gorgonaceans and frondose
macroalgae on hard bottom
Ly)
ee)
Millepora, Palythoa, crustose
corallines, algal turfs, and
Gorgonia
Acropora palmata-reefs
A. cervicornis-reefs
Frondose macroalgae on coarse
sand and rubble bottoms (back
reef)
Beachrock
Montastraea spp.-reefs
99
A. palmata-reefs
A. cervicornis-reefs
Psammophytic algae on
bioturbated sand-plains
39
99
Coralline algal ridges, Millepora,
Gorgonaceans and A. palmata-
reefs
Mixed and scattered corals with
some sand grooves
99
indicate the depth range,
7
marine habitats and
GEOMORPHOLOGICAL
ZONES
Windward fore-reef terrace
bb)
bb)
Continuous peripheral reef
(“Barrier reef”)
Lagoonal terrace
99
Lagoonal basin
99
Discontinuous peripheral
reefs
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99
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ATOLL RESEARCH BULLETIN
NO. 449
SCIENTIFIC STUDIES ON DRY TORTUGAS NATIONAL PARK:
AN ANNOTATED BIBLIOGRAPHY
BY
T.W. SCHMIDT AND L. PIKULA
ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
OCTOBER 1997
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VOIYOLA HANDS NI-WYVd TYNOLLVN SYORIYOL AYA 40 NOLLVOOT
= Wee : wae s eee eee
SCIENTIFIC STUDIES ON DRY TORTUGAS NATIONAL PARK: AN
ANNOTATED BIBLIOGRAPHY
BY
Thomas W. Schmidt! and Linda Pikula”
ABSTRACT
Dry Tortugas National Park, located 110 km west of Key West, Florida, is an elliptical,
atoll-like, coral reef formation, approximately 27 km long and 12 km wide with shallow
water depths ranging from 12-20 m in channels between reefs. In 1935, the area was
designated Fort Jefferson National Monument, the World’s first underwater National
Park unit. Central to the area is Fort Jefferson, America’s largest coastal nineteenth
century masonry fort. In 1992 it was re-designated Dry Tortugas National Park.
Because of the islands’ unique location, the first tropical marine biological laboratory in
the Western Hemisphere was established on Loggerhead Key by the Carnegie Institution
of Washington, Washington, D. C. Following the closure of the Tortugas Laboratory in
1939, aperiodic marine biological assessments have been conducted in response to man-
made and natural environmental perturbations. This annotated bibliography is an
attempt to provide researchers and resource managers with access to the rapidly
accumulating body of information on the park’s natural resources. A total of 424
references (published and unpublished) on scientific studies in, (and what later became)
Dry Tortugas National Park were annotated and indexed according to major scientific
topics. Studies from a wider area were included if they also sampled in Dry Tortugas
National Park.
BACKGROUND
Seven small islands composed of coral reefs and sand in the eastern Gulf of Mexico,
approximately 110 km west of Key West, Florida comprise Dry Tortugas National Park
(Fig.1). The Tortugas, an area known for its bird and marine life and shipwrecks, are an
elliptical, atoll-like, coral reef formation, approximately 27 km long and 12 km wide
with water depths ranging from 12-20-m in channels between reefs.
1 South Florida Natural Resources Center, Everglades National Park, 40001 State
Road 9336, Homestead, Florida 33034, U.S.A.
2 NOAA Regional Library, 4301 Rickenbacker Causeway, Miami, Florida 33149,
U.S.A.
Manuscript received 2 September 1997; revised 2 October 1997
ee ee eee ey ae a ee
II
The Dry Tortugas, discovered by the Spanish explorer Ponce de Leon in 1513 and named
The Turtles, Las Tortugas, were soon read on early nautical charts as "Dry Tortugas" to
indicate they lacked fresh water. Central to the area and located on Garden Key is Fort
Jefferson, America's largest coastal nineteenth century masonry fort. Work was begun in
1846 and continued for thirty years but was never finished. As part of the United States
coastal fortification buildup after the War of 1812, Fort Jefferson was considered critical
for protecting Gulf trade and ports (Murphy, 1993).
Following the Fort's use as a military prison during the Civil War (where the infamous
Dr. Mudd was imprisoned after President Abraham Lincoln's assassination in 1865), and
its abandonment by the Army in 1874, the area was proclaimed a wildlife refuge in 1908,
to protect sooty tern rookeries from egg collectors. In 1935, the area was designated Fort
Jefferson National Monument, the World's first underwater National Park unit. In 1992 it
was redesigned Dry Tortugas National Park to preserve and protect both historical and
natural features.
Early descriptive observers of Dry Tortugas natural resources include Louis and
Alexander Agassiz during the 1850's, and the research vessel Blake in 1877 and 1878.
Their visits resulted in a detailed map of the islands, and a description of benthic marine
communities by Agassiz in 1888.
In 1903, Alfred G. Mayer, under the auspices of the Carnegie Institution of Washington,
recommended that a tropical marine biological research laboratory be established at the
Tortugas (as opposed to other Caribbean sites) because of their isolation from
continental land masses, lack of commercial fisheries, lush reefs, clear waters and
proximity to the Gulf Stream. In 1904, Mayer selected Loggerhead Key as the site for
Carnegie's Tortugas Marine Laboratory, the first tropical marine laboratory in the
Western Hemisphere (Fig.2). Following the closure of the Laboratory in 1939, relatively
few investigations were conducted in the Tortugas until the National Park Service (NPS)
began in 1975, a series of cooperative, bench-mark studies to evaluate long-term changes
in Marine resources in combination with the earlier Carnegie Laboratory studies. Since
the initial Tortugas Reef Atoll Continuing Transect Studies (TRACTS) work of 1975-76,
aperiodic biological assessments have been conducted in response to man-made and
natural environmental perturbations.
PURPOSE
The primary purpose of this annotated report is to provide researchers and resource
managers with a readily accessible document on the rapidly accumulating body of
information on the natural resources of the Dry Tortugas. With the recent
implementation of the Florida Keys National Marine Sanctuary, adjacent to the Park's
boundary, there is a dire need for a scientific database that is centrally located, coherently
organized, and directly related to the future and ongoing management and regulation of
marine resource activities.
Ill
No complete bibliography of the scientific studies on the park's marine and terrestrial
natural resources has been undertaken. In this report we have attempted to list published
and unpublished reports from many fields which we feel will be useful as a starting point
for natural science studies to be conducted at the Dry Tortugas for decades to come (Figs.2&3).
METHODOLOGY
The present bibliographic database containing 424 references was compiled using
PROCITE software, and covers the period of approximately 1878-1996, with the
exception of one report dated 1820. Most 1996 papers were listed through August.
Arrangement is alphabetical by senior author and title. Entries are numbered in
sequence, and each includes a complete bibliographic citation with abstract or summary
While some attempt has been made to achieve uniformity in style, in many cases the
terminology, spelling, capitalization, and phraseology of the original author or abstractor
have been retained. Abstracts obtained from the Carnegie Institution of Washington's
publication citations were adapted from the author's summaries of results. This
bibliography includes books, book chapters, scientific articles, theses and dissertations,
workshop and conference proceedings, reports, and government publications. No attempt
was made to include articles from newspapers or popular boating or sport magazines
Several maps and charts are cited, however.
Research citations were indexed by broad fields of study, specialty sub-headings, and by
both senior and joint authors. Each citation is listed under as many subject headings as
is appropriate for the cited article. This cross indexing system was constructed using
PROCITE.
The geographic boundaries for citations in this bibliography are that the work was done
either completely or partially within the Park boundary, which is the 60' contour line
(Fig.4). Recently, a few studies were undertaken adjacent to or on the Park boundary,
they were included also. Although all Carnegie Institution of Washington published
studies were included if they were conducted at the Dry Tortugas, those studies that were
conducted and identified as solely in the "Gulf Stream" or at satellite marine laboratories
in Jamaica, Trinidad, Puerto Rico, Bahamas, or in the tropical Pacific were not included
A broad range of marine and terrestrial topics were found, including vegetation, marine
algae, invertebrates, sea water composition, and geology. The major topics were sub-
divided into those specialty areas that are shared most often among the studies examined.
For example, sea water composition was sub-divided into salinity and temperature, the
parameters measured most often. In many cases, inclusion or exclusion of a given
reference within a major topic area or specialty sub-heading was a subjective decision.
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LITERATURE SEARCHED
The bibliographic search was predominantly done at South Florida Natural Resources
Center Library, Everglades National Park, Homestead, Florida, and at the NOAA
Regional Library, Miami, Florida. No starting date was established for the references in
this compilation. The senior author began assembling marine archival materials (e.g.,
raw data sheets, correspondence, maps, etc.) and published and unpublished research
results from major scientific studies conducted by NPS scientists and contractors
working at the Tortugas. Pertinent record files were also searched at Dry Tortugas
National Park
The Park Library contains a complete 35 volume set of the Carnegie Institution of
Washington's Tortugas Laboratory Papers. Volumes 1-6 were titled "Papers from the
Tortugas Laboratory of the Carnegie Institution of Washington" (1908-1914), while
volumes 7-14 were "Papers from the Department of Marine Biology of the Carnegie
Institution of Washington" (1915-1926), Volumes 15-29 were issued as "Papers from
Tortugas laboratory of Carnegie Institution of Washington" (1928-1936) and volumes
30-35 were titled "Papers from Tortugas Laboratory" (1936-1942). Each volume was
given a separate publication number by the Carnegie Institution.
The Year Book Series of the Carnegie Institution, which contains annual summaries by
individual investigators on observations and results obtained during their visits to the
Tortugas, were searched at the University of Miami's Richter Library, Coral Gables,
Florida. The Richter Library contains volumes 1-12, and 20-32. Copies of annual
investigator reports for volumes 13-19 and 33-39 were obtained from the Carnegie
Institute of Washington, Washington, D.C. In nearly all cases it was found that the
principal investigators published their final evaluations and conclusions in "Papers",
while the Year Book contained mostly duplicative, preliminary, or unsubstantiated
observations. For these reasons, and due to time constraints, we decided to cite only
Year Book contributions for investigators who did not complete and publish their
conclusions in the "Papers" series. For example, W. H. Longley published in both
series, but is only cited in this bibliography under "Papers" (however, his Year Book
citations can be found in the "literature cited" section of his contributions to "Papers".
S. Yamanouchi, however, only published in the Year Book, and is cited here as such.
We searched documents regarding the Tortugas Laboratory during two visits to the
Carnegie Institution in Washington D.C. Work conducted at the Tortugas Laboratory
has been published in a wide range of journals. For example, a list of scientific writings
produced by activities at the Laboratory during Mayer's directorship can be found in
Papers Tortugas Laboratory 19:80-90. Many publications continued to appear in the
literature following the closure of the Laboratory in 1939.
On-line database searches were conducted during 1993-96 at the NOAA Miami Regional
Library. Subject index terms such as coral reef, geology, vegetation, marine algae, fish,
Vili
etc., were used to search on a variety of DIALOG electronic databases including the
following: BIOSIS PREVIEWS, Dissertation Abstracts, Oceanic Abstracts,
SCISEARCH, Ei Compendex, INSPEC, and GEOBASE. These individual CD’s
were also searched: Aquatic Sciences and Fisheries Abstracts, Life Sciences Collection,
Earth Sciences, GeoRef, NTIS (National Technical Information Service), GPO
(Government Publications Office), and the OCLC (Online Computer Library Center) on-
line catalog.
Pertinent theses, dissertations, and journals identified in the abstracted literature were
obtained via interlibrary loans.
This current compilation undoubtedly does not list all available literature that might be
useful in conducting research, monitoring, and resource management of the park's natural
resources. There may be as many as 100 additional scientific papers generated from the
Carnegie era. We would greatly appreciate additional references to the Tortugas
literature, and if a sufficient number of additional articles become available, we will
produce an addendum to this report.
ACKNOWLEDGMENTS
This report has benefited from the help of many people over the past 6 years. The
original project was prompted and supported by Dr. Michael Soukup during his tenure as
Director of the South Florida Natural Resources Center. Wayne Landrum, Facility
Manager and Carolyn Brown-Wiley, Chief Ranger at Dry Tortugas National Park
provided logistical support and took special interest in providing guidance to the
pertinent files at Fort Jefferson. Ray Bowers, John Strom, and Pat Craig of the Carnegie
Institution in Washington, D.C. permitted us to search their Tortugas Laboratory files,
assisted in duplicating activities, and provided insightful discussions and original
photographs of the Marine Laboratory.
We thank George Stepney and Maria Bello of NOAA’s Regional Library in Miami for
acquiring many interlibrary loans. Special recognition goes to the staff of the South
Florida Natural Resources Center including Marnie Lounsbury for photocopying and
collating much of the Carnegie texts, Barry Wood who produced the map figures, and
Mario Alvarado who expertly produced the author and subject indexes using PROCITE.
Dr. William B. Robertson, Jr., United States Geological Survey/Biological Resources
Division contributed numerous references and provided encouragement during the
earliest stages of the project
Valuable comments were provided by Elaine Collins of the NOAA Central Library,
Silver Spring, MD and Bob Hamre, former technical editor for the US Forest Service,
assigned to the Beard Center under the NPS “Volunteer-in-Parks.” program.
Finally, our thanks to Carol Watts, Chief of the NOAA Libraries and Information
Science Division, Janice Beattie, Chief of NOAA Libraries Public Services Division, Dr.
Tom Armentano, Chief of Biological Resources, South Florida Natural Resources
Center, and Dr. Caroline Rogers, United States Geological Survey/Biological Resources
Division for their financial support and encouragement.
ANNOTATED BIBLIOGRAPHY
1. Agassiz, A.. 1888. The Florida reefs. Three cruises of the United States Coasts and Geodetic
Steamer 'Blake' in the Gulf of Mexico, in the Caribbean Sea and along the coast of the
United States, from 1877 to 1880., V. 1, Chapter 3, pages 52-92. Houghton, Mifflin and
Co. New York. 314 pp.
While the Steamer "Blake" was mostly involved in deep water dredging operations in the
Gulf of Mexico and the Caribbean Sea (1877-78), a five-week visit to Fort Jefferson (Dry
Tortugas) provided the author with an opportunity to work in a laboratory-like situation, to
examine carefully the topography of the different groups of corals characteristic of the
Florida reefs and to give an extended account of the Florida reefs in a special chapter of
this book. The Tortugas, as described by Agassiz, form the most recent of the cluster of
the Florida reefs, and it is here where he begins a topographical sketch of the Florida reefs
from the Tortugas to Cape Florida.
. 1888. The Tortugas and Florida Reefs. Memoirs of the American Academy of Arts and
Sciences, Philadelphia. Centennial ed., V.II, :107-132.
VII entitled "Explorations of the surface fauna of the Gulf Stream, under the auspices of
the United States Coast Survey”.
Agassiz reports on the formation of the Florida Reefs, commenting on the theories of
Darwin, LeConte, and Hunt on this subject. At the time of this article it was believed that
the elevation of the Florida Plateau, from Cape Florida southward to the Dry Tortugas and
the Yucatan Banks, was based on the accumulation of coral sands, as well as animal debris
brought to it by great Atlantic equatorial oceanic currents, the Gulf Stream and prevailing
winds, and at which time, reef-building corals could flourish and a reef would be formed.
They speculated that corals could not thrive below 6 or 7 fathoms, because siltation ooze
would sink to the bottom and choke the corals. Coral reef formations were assumed to be
established near strong equatorial currents which were suppliers of food for the reef-
building corals. It was assumed that corals grow towards the surface as fast as the ooze
deposited has closed up the circulation of the lower levels. At the time of Darwin little was
known of limestone deposits formed by the accumulation of animal decay. Thus
explanations of reef formation other than elevations as a result of submerged mountains
and subsidence were not investigated. The Tortugas Reefs probably are newly developed,
as they have not been above the sea long enough to have received the flora and fauna
characteristic of the Keys north of Key West.
3. Andres, B. A. 1991. Migration of sharp-shinned hawks in the Dry Tortugas, Florida USA. Wilson
Bulletin 103, no. 3: 491-93.
Some species of hawks have been found to make long water crossings during migration.
One of the species, the sharp-shinned hawk Accipiter striatus, seldom undertakes water
crossings of >125 km. However, large numbers of sharp-shinned hawks are observed
every fall in the Florida Keys including the Dry Tortugas, where water crossings are quite
common. No information, however has been gathered concerning their migration after
reaching the Tortugas. Based on wind speed, wind direction, and binocular observations
made on Garden Key of six species of hawks (sharp-shinned hawks dominated the
observations), it was found that sharp-shinned hawks are deliberately initiating an over-
flight across the Gulf of Mexico directly to Central America.
4. Austin, O. L. Jr., W. B. Robertson Jr. and G. E. Woolfenden. 1972. Mass hatching failure in Dry
Tortugas sooty terns, Sterna fuscata. Proceedings of the International Ornithological
Congress. 627. Netherlands.
The author attributes a mass hatching failure among 50,000 pairs of sooty terns (Sterna
fuscata) nesting on the Dry Tortugas to damage caused by sonic booms from low-flying
military aircraft. Theoretically, eggshells and embryonic tissues should withstand pressures
much greater than those generated by even the most intense sonic booms.
5. Bailey, E., G. E. Woolfenden and W. B. Robertson Jr. 1987. Abrasion and loss of bands from Dry
Tortugas sooty terns. Journal of Field Ornithology 58, no. 4: 413-24.
During the past 25 years more than 400,000 sooty terns (Sterna fuscata) have been banded
at Dry Tortugas, Florida, with size 3 aluminum bands of several different alloys. Based on
large samples of bands removed from the terns, regression lines were established for each
of four alloys. Differences in the slopes of the regression lines for certain of the four alloys
demonstrated differences in rates of abrasion. Band loss was evident for bands of the
fastest abrading alloy (2-SO) that were carried more than 20 yrs by terns banded as chicks
because all band weights fell above an extension of the regression line. For this alloy, the
plots of weight loss showed that band loss becomes significant at 86% of original weight.
Bands of alloy 2-SO began reaching 86% of original weight at age 14 when placed on
adults and 20 when placed on chicks. The regression lines for the other 3 alloys suggest
that loss is likely after 17-28 yrs for bands placed on adults and after 20-25 years for bands
placed on chicks. Band loss probably occurs through abrasion of the inner surface, which
increases the inner diameter until the band can slip over the toes. Any gap that develops
would hasten loss.
6. Bailey, P. L. 1938. Regeneration in sabellids. Carnegie Institution of Washington, Year Book 37:
84-85.
Sabellid worms collected in the moat on Garden Key proved to be suitable for fixation
techniques needed to conduct various regeneration experiments to determine the effects of
chemical solutions on the cells.
7. Baker, B. 1994. Partitioning the National Marine Sanctuary. Bioscience 44, no. 7: 497.
A management proposal is described to establish five zones in the Florida Keys National
Marine Sanctuary: replenishment reserves, sanctuary preserve areas, research-only zones,
wildlife management zones, and special-use zones. The Sanctuary encircles the Florida
Keys, including the Dry Tortugas, for 2800 square nautical miles.
8. Ball, S.C. 1918. Migration of insects to Rebecca Shoal Light-Station and the Tortugas Islands, with
special reference to mosquitoes and flies. Papers Tortugas Laboratory 12: 193-212.
Carnegie Institution of Washington Publication Number 252.
The circumstances which suggested the desirability of such investigations were the
repeated experiences of Dr. Mayer and other scientists at Tortugas, Florida, in connection
with the occurrence there of mosquitoes. These insects were abundant on Loggerhead Key
only after northerly winds of several hours' duration, under conditions favorable to their
migration from the mainland of Florida. Rebecca Shoal light-station was chosen as the
study site, because of its isolation from the mainland and other keys, and because of its
freedom from all except easily controllable breeding-places for mosquitoes. It was found
that large numbers of mosquitoes and house-flies are carried by northerly and southerly
winds to Rebecca Shoal light-station and the Tortugas Islands from Florida and Cuba.
Easterly winds bring a few of these, as well as smaller numbers of blow-flies, horse-flies,
and gnats from islands east on the Florida Reef. Occasionally Odonata, Neuroptera, and
Lepidoptera are carried by the winds to these parts of the reef. Sarcophagidae breed in land
crabs at Tortugas .
9. Ballantine, D. L. 1996. New records of benthic marine algae from Florida. Gulf of Mexico Science
118 i iets),
Seven species of benthic marine algae are newly reported from the Dry Tortugas, Florida.
These are Halimeda hummii Ballantine (Chlorophyta), Audouinella ophioglossa
Schneider, Botryocladia uynnei Ballantine, Champia viellardii Kutzing, Monosporus
indicus Borgesen, Hypoglossum rhizophorum Ballantine et Wynne, and Rhodogorgon
ramosissima Norris et Bucher (Rhodophyta). Monosporus indicus is reported for the first
time from the Atlantic Ocean. The Dry Tortugas represents the northern distributional
limit for the remaining species reported, except Audouinella ophioglossa and Botryocladia
uynnel.
10. Bartsch, P. 1919. "The bird rookeries of the Tortugas." Smithsonian Institution Annual Report for
1917, 2512. Smithsonian Museum.
The author states that the most interesting island of the Tortugas group is Bird Key (circa
1908). Of the 32,810 birds listed for the islands, 31,200 center about that Key. A
numerical listing of the summer birds is given. These rookeries were first brought to the
attention of ornithologists by John Audubon in his ornithological biographies. He gives an
account of a visit in May 1832. A first list of birds observed in the Dry Tortugas is given
by W.E.D. Scott in his paper on birds observed during parts of March and April 1890.
Drs. John B. Watson and K.S. Lashley of Johns Hopkins University made an extensive
study of the wild bird colonies there, hoping to throw light on the homing instinct. The
article ends with an extensive listing of bird sightings in the Tortugas up to 1919.
11. ————. 1920. Experiments in the breeding of Cerions. Papers Tortugas Laboratory 14: 1-55.
Carnegie Institution of Washington Publication Number 282.
Breeding experiments were conducted to determine if various forms of Cerion colonies
were fixed forms, that is, will generations yield the same mode in measurement, or will
changes in the local environment from season to season affect the developing organisms to
such an extent as to produce an unending series of slight variations? Introduced forms
were placed where native species existed. Colonies of these land snails were planted on
Keys in the Dry Tortugas in 1914, 800 in 1915 on Loggerhead Key, and a third planting in
1916 of 8,317 specimens. A comparative anatomical discussion of the five species of
Cerion involved in the breeding experiments is given.
12. ————. 1915. Report on the Bahama Cerions planted on the Florida Keys. Papers Tortugas
Laboratory 8: 203-12.
Carnegie Institution of Washington Publication Number 212.
A study is made of the two races of Bahama Cerion transplanted to the Florida Keys in
1912. The conditions of the Cerion colonies are described.. Illustrations show the extent of
the changes between the first generation and the parent generation. Changes that have taken
place in the second generation in shell, color and sculpture are discussed.
13. . 1919. Results in Cerion breeding. Proceedings of the Biological Society of Washington, 32.
Journal Washington Academy of Science 9:657 (abstr.)
A short account is given of Dr. Paul Bartsch's report on the breeding of Cerions
transplanted from Andros Island in the Bahamas to the Dry Tortugas.
14. ———. 1916. Visit to the Cerion colonies in Florida. Smithsonian Explorations 66, no. 17: 41-44.
The author visited the Dry Tortugas through the auspices of the Carnegie Institution and
the U.S. National Museum to observe the transplanted Bahamian Cerion colonies. He
reported finding many adult specimens of the first Florida generation. No adult second
generation specimens were found. Four hybrid specimens between the native Cerion
incanum and the transplanted Bahama stock were obtained.
15.
16.
17.
18.
ID)
20.
Bellow, T. and C. Winegarner. 1975. Nesting of brown pelicans Pelicanus occidentalis on the Dry
Tortugas, Florida. Florida Field Naturalist 3, no. 2: 47-48.
On 14 June 1974 on Bush Key, Dry Tortugas, Florida T. Bellow and C. Winegarner found
5 Brown Pelican nests about 12 feet above ground in the white mangroves (Laguncularia
racemosa) along the north shore. Nineteenth-century records of pelicans breeding on the
Dry Tortugas are ambiguous. It appears that a few pairs did breed on the Tortugas in the
mid-1800's, but by late in the century none did so. This record is the first reported nesting
of this species in the 20th century on these ornithologically well-known islands. Three of
the nests found in 1974 contained 2 eggs each, one nest was empty, and the fifth was not
checked.
Bellow, T. H. 1979. A cardinal at the Dry Tortugas, Florida. Florida Field Naturalist 7, no. 2: 31.
The southern range of the nonmigratory cardinal (Cardinalis cardinalis) extends through
the Florida Keys, but is considered rare in Key West. An April observation of a cardinal
on Garden Key represents the second published record of this species at the Tortugas.
Bennett, F. M. Commander. 1909. A tragedy of migration. Bird-Lore 11: 110-113.
On, April 14, 1909 a violent storm hit the Florida Keys, including the Dry Tortugas. An
apparent bird migration was in progress at the time of the storm. On April 20th the author
went to the Dry Tortugas and observed hundreds of dead birds, and tens of thousands of
injured and exhausted birds. A listing of the types of birds observed is given.
Berrill, N. J. 1938. Budding in polystryelid ascidians. Carnegie Institution of Washington, Year
Book 37: 85.
The area of the budding rudiment relative to the size of the parent zooid was closely related
to the size of the parent zooid, and the general nature of the colony.
Blinks, L. R. 1926-1929. Electrical conductivity in Valonia. Carnegie Institution of Washington,
Year Book.
Note published as follows: 1926, v. 25, p. 240; 1927, v. 26, p. 217-18; 1928, v. 27, p. 270-
71; 1929, v. 28, p. 280.
This study makes use of the good supply of Valonia at Tortugas for studies on the
variability of electrical resistance in protoplasm. Causes of uncertainty are discussed.
Bohnsack, J. A., D. E. Harper and D. B. McClellan. 1994. Fisheries trends from Monroe County,
Florida. Bulletin of Marine Science 54, no. 3: 982-1018.
Fishing is an important activity in the Florida Keys National Marine Sanctuary (FKNMS).
Concern exists that excessive fishing could be deleterious to individual species, disrupt
marine ecosystems, and damage the overall economy of the Florida Keys. We examined
data from commercial, recreational, and marine life fisheries in Monroe County, Florida.
Invertebrates comprised the majority of commercial landings. In 1992, the total reported
commercial landings were composed of 54% invertebrates (4.09 x 10 kg) 28% reef fishes
(2.19 x 10 kg), and 21% non-reef fishes (1162 x 10 kg). In the recreational headboat
fishery, reef fishes accounted for 92% of 0.107 x 10 kg average total annual landings from
the Dry Tortugas and 86% of 0.201 x 10 kg landed from the Florida Keys since 1981.
Average annual landings for other recreational fisheries were estimated at 1.79 x 10 kg for
reef fishes (45%) and 2.17 x 10 kg for non-reef fishes (55%) from 1980 through 1992.
Estimated landings from the Dry Tortugas did not show distinct trends and were highly
variable. Finer resolution of catch and effort data are needed, especially for recreational
fisheries. Landings for some species varied greatly over time. The most conspicuous
declines were for pink shrimp, combined grouper, and king mackerel, while the most
conspicuous increases were for amberjack, stone crab, blue crab, and yellowtail snapper.
Landings of spiny lobster have remained constant. Fisheries closed to harvest included
queen conch, Nassau grouper, jewfish, and stony corals. Effective fishing effort has
increased over time with more participants and more effective fishing technology. Since
1965, the number of registered private recreational vessels has increased over six times,
while the number of commercial and headboat vessels has remained stable. The number of
management actions have continually increased and become more restrictive with increased
fishing effort. Comparison of fisheries was complicated because different fisheries
targeted different species and different sized organisms. Also, landings were sometimes
reported by numbers and sometimes by weight. Measures of reproductive value and
spawning potential are suggested as useful parameters for comparing effects of different
fisheries. The new FKNMS provides a unique opportunity to shift management emphasis
from a species approach to an ecosystem and habitat based approach.
21. Bortone, S. A., P. Rebenack and D. M. Siegel. 1981. A comparative study of Diplectrum formosum
and D. bittatum (Pisces: Serranidae). Florida Scientist 44, no. 2: 97-103.
Specimens of the simultaneously hermaphroditic fish species Diplectrum formosum, the
sandperch, and D. bivittatum , the dwarf sandperch, were collected near the Dry Tortugas,
Florida, by means of shrimp trawl during December 1976. Stomach contents of 326 D.
formosum (100 empty) and 325 D. bivittatum (131 empty) revealed little or no differences
in their food habits relative to number and volume of food items, size of food items or the
contribution, in grams each food item makes to each fish. Both species primarily
consumed amphipods, shrimp, crabs, fish, and polychaetes. Temporally, both species fed
at the same 2 diurnal periods. Species were collected sympatrically but there were areas
where each species dominated in relative abundance.
22. Boschma, H. 1929. On the postlarval development of the Coral Maeandra aerolata (L.). Papers
Tortugas Laboratory 26: 129-47.
Carnegie Institution of Washington Publication Number 391.
During six weeks in July and August 1925, the author studied Maeandra areolata for
researches on the food of reef-corals at the Carnegie Laboratory in the Tortugas. Many of
the colonies contained ripe larvae and the author reared these for the study of their
development. The author concludes the development of the endotentacles which appear
constantly in two successive groups of three, resembles in some way the facts recorded by
de Lacase Duthiers (1872) in Actinia mesembryanthemum, some stages of which show a
marked tri-radial arrangement of the tentacles. The data in the literature on the
development of other coral polyps seem to prove that this successive development of the
endotentacles in two groups is an exceptional case. The bilateral arrangement of the septa
in the oldest stages is in accordance with that found by Duerden (1904) in Siderastrea. As
in the majority of corals in which the young stages are known, the septa in Maeandra
develop in two cycles, first the six endosepta and soon afterward the six exosepta.
23. Bowles, A. E., F. T. Awbrey and J. R. Jehl. 1991. Effects of high-amplitude impulsive noise on
hatching success: a reanalysis of the sooty tern incident, HSD-TR-91-0006. BBN
Laboratories, Inc., Canoga Park, California.
This article attempts to refute the Austin article which attributed a mass hatching failure
among 50,000 pairs of sooty terns (Sterna fuscata), who had nested on the Dry Tortugas to
sonic boom damage from military aircraft. Theoretically, eggshells and embryonic tissues
should withstand pressures much greater than those generated by even the most intense
sonic booms. An experiment was conducted to test whether impulsive noise could be
responsible for the hatching failure. Four pest control devices were exploded near chick
eggs in various states of development: 20 chicken and 20 quail eggs. The mean peak flat
sound pressure level 177.3 db re 20 upa; mean CSEL of 139; mean frequency 620 Hz. No
cracking damage similar to that of the Dry Tortugas eggs occurred. Hatch rates and
weights between control and exposed embryos were not significantly different.
6
24. Bowman, H. H. M. 1918. Botanical ecology of the Dry Tortugas. Papers Tortugas Laboratory 12:
109-38.
Carnegie Institution of Washington Publication Number 252.
As the name of these islands indicates, their vegetation is characteristically xerophytic,
although the rainfall is sufficient to assure the plants the necessary amount of water. The
plants are very interesting when a close study is made of their individual characteristics.
The opportunity for such study was given the writer during the summers of 1915 and 1916,
while pursuing another line of botanical research at Loggerhead Key, where a marine
laboratory is maintained. The Tortugas are really the westernmost of all the Florida Keys,
but are more detached from them and have different geological and botanical aspects.
Species distributional maps were created for each of the eight islands of the Tortugas Atoll.
In this treatment of the species in the Tortugas it has been aimed to give some idea of the
character of the dry-climate plants inhabiting these islands, their distribution, and
particularly the changes which have occurred on the various keys since Lansing's 1904
survey, with an attempt to analyze the reasons for such changes. Notes on the marine
ecology of the Tortugas also are presented, along with descriptions of dominant
submergent vegetation.
25. Boyden, A. 1934-1939. Serological study of the relationships of some common invertebrata.
Carnegie Institution of Washington, Year Book.
Note: published as follows: 1934, v. 33, p. 248-49; 1936, v. 35, p. 82; 1939, v. 38, p. 218.
Preliminary results obtained through the study of the antigens collected from various major
groups of animals at Tortugas were summarized. Blood relationships within Mollusca and
Crustacea were emphasized.
26. Bradbury, R. C. 1992. First Florida record of variegated flycatcher Empidonomus-varius at Garden
Key, Dry Tortugas. Florida Field Naturalist 20, no. 2: 42-44.
The variegated flycatcher occurs throughout most of South America east of the Andes. The
species migrates northward between September and February after breeding in the middle
and southern part of the continent. It winters in the Guianas, northern Brazil, Venezuela,
Colombia, and eastern Peru. This article describes observations of a variegated flycatcher
in Florida, representing the first record in Florida and the third in the United States.
27. Breder, C. M. Jr. 1934. On the habitats and development of certain Atlantic Synentognathi. Papers
Tortugas Laboratory 28: 1-35 (issued Dec. 1932).
Carnegie Institution of Washington Publication Number 413.
In this paper data presented are intended to form a basis for further inquiry into the
comparative development and life habits of the Synentognathi, which includes the familiar
Belonidae (needlefish), Hemiramphidae (halfbeaks), and Exocoetidae (flying fish). The
data on which the present paper are based represent some field studies and laboratory work
on material gathered in the Dry Tortugas, Florida during May and June 1929 The feeding
habits, leaping, and flight during the presence and absence of light, eye specialization,
enemies, and ontogeny and phylogeny are discussed. A new species, Strongylura longleyi
is described. A key to the Tortugas Synentognathi is provided, along with tables, beak
measurement and eye development. The Exocoetidae form the major item of diet of a
variety of sea birds, about the Tortugas at least. Young Exocoetidae pass the most
dangerous part of their day when the sun is low, at which time they are unable to see their
predacious enemies coming from below because of light conditions. The eyes of Belonidae
are provided with elaborate equipment to protect them from the brilliance of their
environment.
28. Breder, C. M. Jr. and J. E. Harris. 1936. Effect of light on orientation and stability of young
plectognath fish. Papers Tortugas Laboratory 29: 23-36 (issued Nov. 1935).
Carnegie Institution of Washington Publication Number 452.
Under certain circumstances some plectognath species will respond to a strong beam of
light by violent gyrations. This was first observed by Breder (1929) at the Tortugas
Laboratory. The mechanism by which these movements are effected, their relationship to
the intensity and duration of the stimulus, and the disappearance of the phenomenon with
advancing age of the animal give rise to a number of interesting problems concerning the
action of the receptor-effector system in these fish. This paper is an attempt to explain this
feature of fish behavior. It was found that small specimens of Monocanthus and
Lactophrys, if exposed to a beam of light, frequently exhibit somersaulting or rotational
movements of great rapidity. Somersaulting is produced by passing the locomotor waves
in opposite direction along the dorsal and anal fins. The rotational movement is
accompanied by the deflection of the dorsal and anal fins to the opposite sides of the body,
the direction of motion of the undulations being usually antero-posterior in both fins.
Various combinations of fin and tail movements may occasionally give other twisting
gyratory movements. The primary response to light is always an attempt at reorientation of
the animal so that the light is incident upon the dorsal surface. In fish kept in complete
darkness and "sensitized" by repeated stimuli, gyrations may continue after the light has
been removed, and even mechanical stimuli may initiate similar paroxysms, the
equilibrating system apparently being more or less permanently deranged. Specimens of
Monocanthus over 50 mm. in length do not usually display this behavior, and species other
than plectognaths show it very feebly or not at all. The integration of gravitational stimuli
into the behavior pattern is apparently not perfectly attained until a comparatively late stage
in development, and light is the primary orienting factor. The gyrations are apparently due
to instability, consequent upon overcorrection.
29. Brinley, F. J. 1937-1938. Studies on the implantation of embryonic fish tissue, with notes on the
spawning habits and development of four species of fish. Carnegie Institution of
Washington, Year Book.
Note: published as follows: 1937, v. 36, p. 86; 1938, v. 37, p. 86-7.
Livers and spleens were transplanted from embryos of the hard head shiner to other
embryos of the same age. No apparent effect on the host was noticed. Eggs of
Pomacentrus and parrot fish were collected for observation, along with nurse shark
embryos. Additional work was performed on the origin of muscular movement in these
species.
30. Brooks, H. K. 1962. Reefs and bioclastic sediments of the Dry Tortugas (abs.). Geological Society
of America. Special Paper 73: 1-2.
Many miscomprehensions exist relative to origin of the Florida Reef track and, in
particular, its southwestern extremity-the banks, shoals, and reefs known as the Dry
Tortugas. They are not an atoll as stated by Vaughan (1914). The component
physiographic features rise from a shallow limestone platform 80 to 100 feet below sea
level. Relief features are banks and shoals of bioclastic sands. Their genesis and
circulation distribution are related to the prevailing seasonal storm patterns. Large patches
of Acropora cervicornus (Lamark) are widely distributed through the area in water less
than 60 feet deep. Live coral on these patches is sparse. Proliferation of the staghorn
corals is slow, but cumulative growth has produced a magnitude of skeletal remains. The
coralla are preserved and are ultimately indurated into a porous rocky mass by the luxuriant
growth of Lithohamnion and its cognate encrusting associates. The shallow reefs of
Garden and Loggerhead Keys, populated by calcareous algae, alcyonarians, and
scleractinians, etc., originate upon a foundation of the remains of these organisms. This
can be seen where erosion in surge channels has exposed the underlying materials.
SIE
a2
33%
34.
Brooks, W. K. 1908. Salpa floridian (Apstein) Part II in the Pelagic Tunicata of the Gulf Stream.
Papers Tortugas Laboratory 1: 75-89.
Carnegie Institution of Washington Publication Number 102.
This rare Salpa about which little is known, has been noted in this paper. Mature
specimens of both stages of Salpa were found, in May 1906, on the surface in the vicinity
of the Marine Biological Laboratory at Tortugas, Florida; and an opportunity was afforded
to study and sketch them while alive, and thus to make additions to, and some slight-
corrections of, the count of the species..
Brooks, W. K and C. Kellner. 1908. On Oikopleura tortugensis, n.sp. a new appendicularian from
the Tortugas, with notes on its embryology in Part IV, The Pelagic Tunicata of the Gulf
Stream. Papers Tortugas Laboratory 1: 73-95.
Carnegie Institution of Washington Publication Number 102.
This species was found in abundance near the Marine Laboratory. The specimens are from
5 to 8 mm. long and occur in great swarms at the depth of 5 to 6 fathoms. A description of
the species is provided.
Brown, D. E. S. 1935. Cellular reactions to high hydrostatic pressures. Carnegie Institution of
Washington, Year Book 34: 76-77.
Physiological studies were carried out on the muscles of crabs and fish collected in deep
(100 fathoms) and shallow water of the Tortugas.
Brown, W. Y. and W. B. Robertson Jr. 1975. Longevity of the brown noddy. Bird-Banding 46, no.
3: 250-251.
Despite its abundance and pantropical range, little published information exists on the
longevity of the brown noddy (Anous stolidus). Woodward (Atoll Research Bull, 164:
280,1972) reported a maximum known survival of 10 years for brown noddies banded as
adults on Kure Atoll , Hawaii. Brown noddies on Manana Island, Oahu, Hawaii (A. s.
pileatus) and the Dry Tortugas, Florida (A. s. stolidus), are among the few populations that
have been banded over a period long enough to provide quantitative data on longevity.
Twelve of the brown noddies banded on Manana before 1948 were recaptured dead or
alive before 1960, the longest interval from banding to recapture being 13 years. On 23
May 1972 Brown recaptured on Manana a brown noddy that had been banded there as a
juvenile on 12 June 1947, 25 years earlier.
35. Bullington, W. E. 1940. Some ciliates from Tortugas. Papers Tortugas Laboratory 32: 179-221
(issued Sept. 1940).
Carnegie Institution of Washington Publication Number 517.
During the summers of 1930, 1931, and 1935, during a special study of spiraling in certain
species of ciliates at Tortugas, there appeared in the author’s cultures from time to time
many other species which seemed to be new or little known. There are now fifteen species,
either new to science or little known, about which it is believed sufficient information is
available to justify their description or redescription. Five of the fifteen species have
previously been described, but none of them is well known. Ten were described as new.
They were characterized by amazing shades of color, yellow and red predominating. The
species here discussed and described constitute only a few of those which have been seen
at Tortugas at one time or another, but these are all the author feels justified in discussing,
at the present time, with the information at hand.
36. Burkenroad, M. 1929. Studies upon plankton and the mechanism of sound production in
Haemulidae. Carnegie Institution of Washington, Year Book 28: 283-90.
Daily tows were made from May 31 to August 19 near Loggerhead Key. The variety of
species and numbers of individuals found disputed the notion that the Tortugas region
"once noted for the variety and richness of its floating life, has gradually become in recent
years an almost desert sea."
37. Caira, J. N. and M. H. Pritchard. 1986. A review of the genus Pedibothrium Linton, 1909
(Tetraphyllidea Onchobothriidae) with a description of two new species and comments on
the related genera, Pachybothrium Baer and Euzet, 1962 and Balanobothrium Hornell,
1912. Journal of Parasitology 72, no. 1: 62-70.
A review of the genus Pedibothrium Linton, 1909 is based on type and voucher specimens.
The type species, Pedibothrium globicephalum Linton, 1909 is redescribed. Descriptions
of Pedibothrium brevispine Linton, 1909 and Pedibothrium longispine Linton, 1909 are
emended. Two new species are described, the generic diagnosis is emended, and a key is
provided.
38. Calder, D. R. 1992. Similarity analysis of hydroid assemblages along a latitudinal gradient in the
Western Atlantic. Canadian Journal of Zoology 70, no. 6: 1078-85.
Shallow-water (0-100 m depth) hydroid faunas reported from 26 locations along the
western North Atlantic coast between the high Canadian Arctic archipelago and the
Caribbean Sea were compared. Species numbers varied widely between locations, but
were highest in the tropics and subtropics, lowest in arctic and subarctic waters, and
intermediate in mid-latitudes. Percentages of species producing free medusae were lowest
in high latitudes, intermediate in low latitudes, and highest in mid-latitudes (especially in
estuaries). In a numerical analysis, similar hydroid faunas were identified at locations (1)
between the high Canadian Arctic islands and the Strait of Belle Isle off western
Newfoundland; (ii) between the Gulf of St. Lawrence and Chesapeake Bay; (iii) between
North Carolina and southeastern Florida (south as far as St. Lucie Inlet), and including the
northern Gulf of Mexico; (iv) in the Caribbean Sea, together with Dry Tortugas and the
oceanic island of Bermuda. The greatest change in hydroid species composition along the
coast appeared to occur around Cape Hatteras.
39. Carrier, J. C., H. L. Pratt Jr. and L. K. Martin. 1994. Group reproductive behaviors in free-living
nurse sharks, Ginglymostoma cirratum. Copeia 3: 646-56.
Mating events of the nurse sharks were observed in a nine-day period in the Dry Tortugas
islands. There were four stages of mating: precoupling, coupling, positioning and
alignment, and insertion and copulation. Films were made of four of the mating events.
Seminal fluid released into the water was obtained following one of the copulations. It
showed the presence of free, nonpackaged sperm cells. Of the fifty mating events observed,
ten of these involved multiple males attempting to copulate with single females.
40. Carrier, J.C. and H. L. Pratt Jr. 1997. Habitat management enclosure of a nurse shark breeding and
nursery grounds. Fisheries Research (In press).
Based on nurse shark breeding studies conducted at Dry Tortugas, a sanctuary for nurse
shark reproductive and nursery activities is being established at Dry Tortugas National
Park.
41. Cary, L. R. 1915. The Alcyonaria as a factor in reef limestone formation. Proceedings of the
National Academy of Science 1: 285-89.
In many areas of the Floridean-Antillean region, Gorgonaceae rather than stony corals
make up the most characteristic feature of the lime-secreting organisms permanently
attached to the bottom. In this paper, data are presented on the amount of material
contributed to reef formation by gorgonians. Three factors were taken into consideration:
spicule content (the amount of lime held as spicules in the colonies), distribution of
gorgonians on the Tortugan reefs (the bulk of the gorgonians on any reef area) and
10
42.
43.
44.
disintegration of the coenenchyma of the colonies and the addition of their spicules to the
reef building materials. Using line surveys and the weight and percentage of spicules in the
colonies, it was found that the amount of lime held as spicules in the tissue of living
gorgonians per acre of reef area is 5.28 tons. Next to the destruction of the colonies by
wave action (storms), the greatest mortality of the colonies is from overgrowth of tissues by
other organisms. The destruction of Tortugan gorgonian colonies was nearly complete in
the hurricane of October 1920. It has been estimated that nearly one-fifth of the gorgonian
colonies are destroyed annually.
. 1918. The Gorgonaceae as a factor in the formation of coral reefs. Papers Tortugas
Laboratory 9: 341-62.
Carnegie Institution of Washington Publication Number 213.
An important constituent of the limestone of coral reefs is the calcium carbonate secreted in
the skeletal structures of Anthozoa and marine calcareous algae. Representatives of the
Hydrozoa were important reef formers in past geological epochs, but in the formation of
modern reefs they constitute a minor factor. Representatives of the Anthozoa, the stony
and flexible corals, are among animals the only important agents in the formation of the
modern reefs. The results of this study show that over large reef areas, in the Tortugas at
least, the gorgonian fauna is by far the most important element contributing to the
formation of reef limestones. The amount of spicules in the tissues of gorgonian colonies
would average at least 5.28 tons to the acre for all of the reefs in the Tortugas group. The
figures given represent only a potential contribution to reef formation but a study of the
normal cycle of changes in the gorgonian fauna of this region has shown that at least a fifth
of this amount of calcium carbonate, as spicules, will be added to the reef limestones
annually .
. 1934. Growth of some tissues of Ptychodera bahamensis in vitro. Papers Tortugas
Laboratory 28: 195-213.
Carnegie Institution of Washington Publication Number 435.
Nearly all refinements and expansions of the technique of tissue culture have taken place
with warm-blooded animals as the experimental material. This line of development has, no
doubt, been followed because of its possible medical application. Technical difficulties
extending this method to invertebrates, where the necessary asepsis is more difficult to
attain, have also played a part. The writer developed a technique which was successfully
applied to some tissues of eleven species of marine animals belonging to seven phyla. In
all cases, both migration and cell multiplication were obtained. Two organisms seemed to
offer particularly favorable material for tissue culture. One was the gastropod Astroea
longispina; the other was the enteropneustan Ptychodera bahamensis. This being the
most convenient material with which to work, investigations in 1932 were confined to the
tissues of this species alone. The technique of a method, using either hexyl-resorcenol or
ultraviolet radiation in amounts harmless to the tissues, for growing in vitro the cells of
marine invertebrates is described. Because of their structure, members of the
Enteropneusta lend themselves especially well to the obtaining of explants composed of
one or of several types of tissue. The growth and reproduction of cells from the caecal
portions of the intestine are recorded in detail. The changes undergone by muscle cells
when removed from the body of an animal show a characteristically reversible series of
stages peculiar to this type of cell. The bearing of the observations on Ptychodera cells to
broader problems of cytology is considered .
. 1915. The influence of the marginal sense organs on functional activity in Cassiopea
xamachana. Proceedings of the National Academy of Science 1: 611-16.
The influence of sense organs (nervous system) on the rate of regeneration was examined
at the Dry Tortugas using the disks of the rhizostomous medusa Cassiopea, which can be
11
separated from the oral arms and kept in dishes of seawater for an indefinite period. Pairs
of disks were examined from which all of the thopalia were removed, while from the other
equal amounts of tissues were removed from the bell margin between the thopalia. In all
instances, the disks where the half on which the thopalia remained regenerated at a more
rapid rate than the inactive half. Other experiments focused on influence of sense organs
on the rate of metabolism as measured by production of carbon dioxide. Carbon dioxide
produced was always greater for the normal disk containing sense organs. It was
concluded by the author that in this type of experiment there is some other form of
metabolic activity which is of greater importance as a source of CO) and which is more
directly under the influence of the sense organs than is the activity of the muscular system.
45. ———. 1916. The influence of the marginal sense organs on the rate of regeneration in Cassiopea
xamachana. Journal of Experimental Zoology 21, no. 1: 1-31.
Studies were conducted on accepting the view of the direct or indirect influence of the
nervous system on regeneration in Cassiopea xamachana collected from the Fort Jefferson
moat at Dry Tortugas. Experiments conducted to determine the influence of sense organs
on the rate of regeneration were inconclusive, when testing entire disks with sense organs
removed, compared with specimens where the sense organs remained because of wide
differences in physiological activity between different individuals. Half disks with sense
organs regenerated more rapidly than those half disks without sense organs. Other
experiments involving electrical stimulus by induction shocks on disk halfs, with and
without sense organs, indicated regeneration is faster in the activated half, than from the
inactive disk. These experiments indicate the rate of regeneration is simply one expression
of the general metabolic activity of an animal, and as such is subject to the influence of the
nerve centers, as are many of the functional activities.
46. 1914. Observations upon the growth-rate and ecology of gorgonians. Papers Tortugas
Laboratory 5: 79-90.
Carnegie Institution of Washington Publication Number 182.
This report provides a record of observations extending over a 3-year period on the growth
rate of Gorgonia flabellum and Plexaura flexuosa on the reefs around the Dry Tortugas,
Florida. Ecological observations are supplemented by observations made in Jamaica. For
effective attachment of the planule, the presence of depressions or cracks into which the
planule could settle appears to be the most important factor. In comparison with young
coral polyps the gorgonian colony has an obvious advantage, in that is most rapid growth is
perpendicular to the surface, which permits its most rapidly growing part to secure food
and oxygen. Wave action during very severe storms is by far the most destructive agent to
which Gorgonia are subjected. It appears that the greatest destruction by storms comes
from the tearing of the Gorgonia colonies from the substrate rather than laceration of
tissue.
47. 1917. Studies on the physiology of the nervous system of Cassiopea xamachana. Papers
Tortugas Laboratory 11: 121-70.
Carnegie Institution of Washington Publication Number 251.
In this paper are gathered the results of several distinct lines of experimentation. They deal
with some phase of the physiology of the nervous system of Cassiopea and represent
portions of a general program of research on the nervous system of the lower animals. On
account of its ability to live under adverse conditions and to withstand practically any type
of operation, Cassiopea is an especially favorable form for experimentation and has been
used as a subject for many researches. The experiments with entire disks, when the rates of
regeneration of specimens on which the sense-organs remained are compared with those of
specimens from which all sense-organs are removed, are inconclusive because of wide
differences in physiological activity between different individuals. When we compare the
12
insulated halves of a disk, on one of which the sense-organs remain, while all of them have
been removed from the other half, it is found that the half-disk with sense-organs always
regenerates most rapidly. When all the sense-organs are removed from a disk and the
halves insulated, the regeneration is faster from the activated than from the inactive half-
disk. These experiments indicate that the rate of regeneration is simply one expression of
the general metabolic activity of an animal, and as such is subject to the influence of the
nerve-centers, as are many other functional activities. Briefly summarized, the results of the
observations made on the starved Cassiopea are as follows: In general the smaller
Cassiopea loses relatively more in weight than does the larger Cassiopea. The percentage
of water found in the entire body is nearly the same in all sizes of Cassiopea. The
nitrogen-content of the entire body is higher in the small than in the larger Cassiopea.
However, the absolute amount of nitrogen found in the starved Cassiopea is considerably
higher than in the normal having the same bodyweight. The loss in weight of the different
parts in the starved Cassiopea remains the same proportionately to those in the normal
Cassiopea.
48.
. 1918. A study of respiration in Alcyonaria. Papers Tortugas Laboratory 12: 185-91.
Carnegie Institution of Washington Publication Number 252.
Although the respiration of many species of invertebrates has been studied, the only
references to that of Alcyonaria are those given by Montuori (1913), who studied two
species, Alcyomeum pallidum and Gorgonia cavolinii. In these experiments the total
weight of the colony was taken as the basis of comparison without taking into account the
proportion of inert skeletal material- the spicules in the first species and the spicules and
chitinous axis in the latter. The observations recorded were made as part of a study of the
ecological factors determining the distribution of Alcyonaria on the coral reefs of southern
Florida. All the species of the genus Gorgonia and the closely related Xiphigorgia, which
have as a group the highest rate of respiration, are next to Briareum the most resistant to
increased temperature. Taken all together these observations indicate that some other
factor is the controlling agency in the ability of a marine organism to withstand high
temperatures. The acidity of the water at the close of the heat experiments was always
greater than in respiration experiments carried on at 27.5° C. This may be only an
expression of the abnormality of their metabolism at high temperatures, or have a causal
relation to the death of the organism.
49. Cate, C. N. 1978. New species of Ovulidae and reinstatement of Margovula pyrulina (A. Adams,
1854) (Gastropoda). Nautilus 92 , no. 4: 160-167.
Eight species of living Ovulidae are described as new, and the species M. pyrulina is
reinstated. The 8 new species are listed as follows: Prionovolva castanea from the Gulf of
Oman; Aperiovula testudiana from Mukaishima, Japan; Primovula santacarolinensis
from Mozambique; P. uvula from Moreton Bay, Queensland, Australia; Crenavolva
periopsis from Java, Indonesia; Speculata advena from off Sand Key, Florida; Cyphoma
rhomba from Fort Lauderdale Reef, Florida; and Psudocyphoma gibbulum from off the
Dry Tortugas Islands, Florida.
50. Chambers, E. L. 1937. The movement of the egg nucleus in relation to the sperm aster in the sea-
urchin, Lytechinus varigeatus. Carnegie Institution of Washington, Year Book 36: 86-87.
With the aid of a camera, 30-second observations on the positions (rate and direction of
movement) of the egg nucleus and sperm aster of Lytechinus were made.
51. Child, C. A. 1992. Shallow-water Pycnogonida of the Gulf of Mexico. Memoirs of the Hourglass
Cruises 9, no. 1: 1-86.
This paper treats 11 species in 8 genera of the Pycnogonida that were collected during the
Hourglass Cruises, a sampling program conducted on the central West Florida Shelf for 28
13
months during 1965-1967. Five benthic stations in depths from 6 to 72 m were sampled
monthly with dredges and trawls among each of two transects. Treatments of 20 more
species in 6 additional genera from other shelf collections are also included to offer a
comprehensive survey of species (a total of 31 species in 14 genera) known from the
continental shelf of the Gulf of Mexico, excluding the Dry Tortugas and the Florida Keys.
Three of these species were previously unreported from the Gulf. Two new species,
Ascorhynchus crenatum and A. horologium, are described from the Hourglass material,
and an additional new species, Anoplodactylus dauphinus, is described from the other
material. Artificial taxonomic keys are provided for all Gulf of Mexico families and
species, and checklists are provided for all species known or expected to occur in the Gulf.
All species are diagnosed and illustrated, and their distributions are given. Only four
species were taken during the Hourglass Cruises with sufficient frequency to allow analysis
of their distributions and abundances.
52. Clapp, R. B. and W. B. Robertson Jr. 1986. Nesting of the masked booby Sula dactylatara on the
Dry Tortugas, Florida. The first record for the contiguous United States. Colonial
Waterbirds 9, no. 1: 113-16.
In both 1984 and 1985 masked boobies (Sula dactylatra) attempted to nest on sandy islets
at the Dry Tortugas, Florida. Nesting attempts failed because the nest sites were washed
away by summer storms. It seems likely that this species will eventually nest there
successfully and will establish a small breeding population. This is the first documented
nesting by this species in the contiguous United States.
53. Clark, H. L. 1919. The distribution of the littoral echinoderms of the West Indies. Papers Tortugas
Laboratory 13: 49-74.
Carnegie Institution of Washington Publication Number 281.
The purpose of this investigation was to determine if the distribution of littoral
echinoderms varied among various northern West Indian islands bounded by Bermuda to
the North, Tobago to the South, and the Tortugas to the west. Five classes of echinoderms
are discussed, including Comatulidea (feather-stars), Asteroidea (sea-stars), Ophiuroidea
(brittle-stars), and Echinoidea (sea urchins). The number of species and numbers of
individuals of the classes are discussed. Of the island areas investigated, the Tortugas
appears to be the richest in the number of species found with 76 littoral echinoderms, with
70 available to collect by hand. The sea urchins, Echinometra lacunter were reported to be
excessively abundant on many reefs, actually occurring by the thousands. The number of
echinoid species found throughout the region and Florida are compared with comments
provided on the origin of echinoids in the West Indies. Next to brittle-stars, holothurians
were considered to be the most abundant of the littoral echinoderms.
54. Clark, L. B. and W.N. Hess. 1942. Swarming of the Atlantic palolo worm, Leodice fucata (Ehlers).
Papers Tortugas Laboratory 33: 21-70 (issued Oct. 1940).
Various organisms show reproductive activity coinciding with the lunar cycle. At the Dry
Tortugas, swarming observations were recorded during 1937-39 on the Atlantic palolo
worm, in association with the quarter-moon phase. Other important factors determining the
time when the worms reproduce, include the maturity of the animal and the amount of
water turbulence. It was concluded that: (1) the stimulating effect of the first-quarter moon
is less than that of the third-quarter, (2) the worms increase in sensitivity to the stimuli
inducing the swarming as they become sexually mature, and (3) wave action and water
turbulence above a certain level induced by an 8 mph wind decreases or prevents swarming
at the Dry Tortugas.
14
a5:
56.
57,
58.
52:
60.
61.
Cole, L. J. 1906. Ant Studies. Carnegie Institution of Washington, Year Book 5: 110.
To investigate the biology of ants at the Tortugas, specimens were collected, and
observations were made.
Collie, M. R. 1979. A Sabine's gull at the Dry Tortugas. Florida Field Naturalist 7, no. 2: 28.
A photographic observation was made by the author during August 1978 of a Sabine's gull
at Garden Key, Dry Tortugas. There are four other records of this species in Florida along
the Atlantic coast, however this sighting represents the first record of the Sabine's gull at
the Tortugas.
Colman, J. 1931. The superficial structure of coral reefs: animal succession on prepared substrata.
Carnegie Institution of Washington, Yearbook 30: 395.
Plant and animal successions were examined on concrete cubes planted in the water at
three sites: Fort Jefferson moat, an iron wreck east of Loggerhead Key, and northwest of
loggerhead Key. Also, a detailed ecological survey of Long and Bush Keys was made.
Conger, P. S. 1924-1935. Diatom studies. Carnegie Institution of Washington, Year Book.
Note: published as: 1924, V. 23, p. 220; 1925, v. 24, p. 221; 1926, v. 25, p. 240; 1927,
v.26, p. 220; 1928, v. 27, p. 271; 1929, v.28, p. 283; 1930, v. 29, p. 323.
Diatom studies were conducted in association with A. Mann. Narrative same as in
reference no. 218.
Conklin, E. G. 1908. The habits and early development of Linerges mercurius. Papers Tortugas
Laboratory 2: 153-70.
Carnegie Institution of Washington Publication Number 103.
The jellyfish, Linerges mercurius, Scyphomedusa, was investigated at Tortugas and Nassau
Harbor. The sudden appearance in great numbers of this species at Tortugas was noted,
followed by their rapid disappearance. Normal movements of the medusae are described
as well. Other phases of early development, including egg-laying, egg structure,
maturation and fertilization, first cleavage, second, and later cleavages, and blastula,
gastula, and planula are described. Experiments on isolation of blastomeres and
centrifugalized eggs are presented. The organization of the egg of Linerges and mechanics
of cell division are described.
. 1908. Two peculiar actinian larvae from the Tortugas, Florida. Papers Tortugas
Laboratory 2: 171-86.
Carnegie Institution of Washington Publication Number 103.
During middle-of-the-day sampling tows at the Tortugas in May 1905, two peculiar larvae
were collected. They did not undergo metamorphasis in aquaria. Natural history notes on
living larvae are provided. Based on literature description, they were probably Zoanthidae
of the Order Hexactinia. A band of strikingly brilliant, locomotor cilia was noted as most
peculiar for these larvae. Their size, shape, and coloration are described. Yellowish-green
symbiotic algae occur in both types and are hypothesized to be associated with their
metabolism and play an important role in their nutrition. The morphology and histological
character of these two types are similar but minor differences are described. Although
these types have only been collected a few times world-wide, they are not considered rare
at Nassau and the Dry Tortugas.
Coonfield, B. R. 1940. Chromatophore reactions of embryos and larvae of Pomacentrus
leucostictus. Papers Tortugas Laboratory 32: 169-78 (issued Sept. 1940).
Carnegie Institution of Washington Publication Number 517.
Interest in the origin of the changing color patterns of fishes and certain other vertebrates,
together with certain questions that have been raised by the work of investigators in their
62.
15
study of this problem in embryos of vertebrates, prompted an investigation of the color
mechanism in developing fishes. Both embryos and larvae of Pomacentrus leucostictus,
which is found in abundance in the Dry Tortugas, were used in this study. It was
concluded that melanin granules migrate within the melanophores of Pomacentrus embryos
as soon as these pigmentary bodies are completely formed. The melanophores of embryos
a few hours of age contract in response to pressure applied with forceps and to a
temperature of about 8°C. The melanophores of a majority of developing embryos, from
their beginning up to a few hours before hatching, are found to be in a stellate state
regardless of whether these young are over a white or a black background or are in total
darkness. A few hours before hatching, the melanophores contract when the embryos are
over a white background, expand when they are over a black background, and contract
when they are in total darkness. This response continues in these young on through the
hatching period and for a few hours after hatching. Larvae of two or more days after
hatching do not show any conclusive response to different backgrounds or to the absence
of light. The eyes of these young fish are believed to have no function in controlling their
melanophore responses. The evidence is in favor of the release of a hormone within the
capsule just before the embryos hatch. This agency either permits or directly causes the
melanophores to respond to various environments.
. 1940. The chromatophore system of larvae of Crangon armillatus. Papers Tortugas
Laboratory 32: 121-26 (issued May 1940).
Carnegie Institution of Washington Publication Number 517.
The ability of certain animals to imitate the color in their background 1s so striking that is
has received the attention of investigators for a considerable period of time. This feature
has been observed principally in fishes, amphibians, and reptiles of the vertebrates, and in
crustaceans of the invertebrates. This paper adds to this field of study the results of
observations on the reactions of the chromatophore system of larvae of Crangon
armillatus. The erythrophores of normal larvae of Crangon armillatus react as follows
according to different backgrounds: the pigment is dispersed when the animals are kept in a
white illuminated bowl. The erythrophores of enucleated specimens show the following
conditions when subjected to different backgrounds: the pigment becomes concentrated
when the specimens are kept over a white illuminated background; the pigment is
dispersed when these specimens are subjected to black illuminated bowls. The time
required for concentration of pigment in the erythrophores is much longer than that
required for its dispersion. Ablation of the eyes permits the erythrophores to react directly
to stimulations caused by different backgrounds .
63. Coutiére, H. 1910. The snapping shrimps (Alpheidae) of the Dry Tortugas, Florida. Proceedings of
the United States National Museum 37: 485-87.
The Alpheidae collected by Dr. McClendon at the Tortugas in 1908 are discussed. The
Alpheidae are referable to eight different forms, including one new species and one new
subspecies: Alpheus formosus Gibbes. Alpheus cristulifrons Rathbun and Alpheus
armillatus H. Milne-Edwards.
64. Cowles, R. P. 1908. Habits, reactions, and associations in Ocypoda arernaria. Papers Tortugas
Laboratory 2: 1-41.
Carnegie Institution of Washington Publication Number 103.
On Loggerhead Key, investigations were made on the behavior of Ocypoda arernaria. It
was found that adult ghost crabs build two kinds of burrows. One consists of a single
tunnel extending down in the sand for 3 to 4 feet. The other is similar, except that it is
shorter and has a passage branching off from it, which is used for escape. Young ocypodas
make short burrows, only a few inches long, which often extend vertically downward.
Breeding in the region of Loggerhead Key probably occurs in the spring and early summer.
16
65.
Ocypoda is a scavenger and a cannibal. The eyes do not seem to play an important role in
the detection of food, but they undoubtedly lead individuals to objects which may be food.
That Ocypoda is stimulated by odors was not conclusively shown, but certain experiments
point strongly in that direction. The eyes are highly developed, so far as crustacean eyes
are concerned; they are quite sensitive to large differences in the intensity of light; they do
not react to different colors; they aid much in the search for food, in the detection of
enemies, and in the accuracy of locomotion. Ghost crabs probably do not have vision such
as that of the human eye, nor do they see the color and finer characters of the surface of an
object, but they undoubtedly see its outlines and possibly some of the more evident
irregularities of the surface made evident by differences in lighting. The color-pattern seen
through the carapace of Ocypoda changes in intensity under different conditions of
temperature and light. In the absence of light when the temperature is anywhere between
22° C. and 45° C., and undoubtedly when it is even lower or higher, a light coloration
occurs. Generally in diffuse light and even direct sunlight a dark coloration appears,
provided the temperature is not too high. Usually at low temperatures, not above 35° C., a
light coloration is the rule, and it occurs independently of the intensity of light. At high
temperatures, above 35° C., a light coloration is the rule, and it occurs independently of the
intensity of light. No indication of audition was observed in Ocypoda. The so-called
"auditory organs” are equilibrating organs. Ocypoda has a stridulating ridge on the palm of
its large chela.
. 1911. Reaction to light and other points in the behavior of the starfish. Papers Tortugas
Laboratory 3: 95-110.
Carnegie Institution of Washington Publication Number 132.
Experiments were designed to test the reactions of starfish to light. Two species were used
Echinaster crassispina and Astropecten duplicatus. Both are migratory and are found in
open waters over sandy bottoms, in areas generally exposed to light. In the Tortugas
laboratory, starfish were placed in aquaria wooden boxes and tested for movement up in
response to light, inclines, vertical walls, and tilted floors. Every specimen reacted
positively, moving toward bright light. Even with eye-spots removed, movements towards
light are positive, but not as quick as in normal individuals. When tested in different
degrees of water temperature the reaction to light was positive at ordinary temperatures.
Quality of light was tested using various color screens (UV, violet, blue, green, yellow,
orange and red) in a box with closed and open ends. The source of light was sunlight. Ina
series of 10 tests with varied orientation and handling, in nearly every test starfish moved
toward light without hesitation.
66. Criales, M. M. and T. N. Lee. 1995. Larval distribution and transport of penaeoid shrimps during
the Tortugas Gyre in May-June 1991. Fishery Bulletin 93: 471-82.
As part of the Southeast Florida and Caribbean Recruitment (SEFCAR) project, penaeoid
shrimp larvae were collected during the spring and summer cruise of the RV Longhorn in
the Lower Florida keys and Dry Tortugas from 29 May to 30 June 1991. Larvae of the
pink shrimp, Penaeus duorarum, and the rock shrimp, Sicyonia sp., were distributed
inshore close to the Dry Tortugas Grounds adjacent to the boundaries of Dry Tortugas
National Park, whereas larvae of the oceanic shrimp Solenocera sp. showed mainly an
offshore distribution. Significant concentrations of Solenocera sp., Sicyonia sp. and P.
duorarum larvae at the Tortugas transect in early June were found within and above the
seasonal thermocline, while the cold cyclonic Tortugas Gyre was intensively developed.
For Solenocera sp., which spawn on the outer ridge of the gyre followed by onshore
Ekman transport. Penaeus duorarum, which spawn in the shallow Tortugas Grounds,
showed a mode of zoea II-III progressing to postlarvae I at the Tortugas Grounds during
the 15 days in which the drifter Halley recirculated in the interior of the Tortugas Gyre.
Retention of P. duorarum larvae by the internal circulation of the gyre at the spawning
1,
grounds may be an important mechanism for local recruitment of these shrimp to the
nursery ground of Florida Bay, Everglades National Park.
67. Cushman, J. A.. 1922. Shallow-water Foraminifera of the Tortugas region. Papers Tortugas
Laboratory 17: 1-85.
Carnegie Institution of Washington Publication Number 311.
The paper gives the results of a study of collections made in the waters about the Tortugas
Laboratory of the Carnegie Institution of Washington. Collecting was done largely from
the boats, the most satisfactory method that was used with the Darwin. Collecting in the
moat at Fort Jefferson in shallow water on Long Key, as well as on the reefs and flats, was
done by hand. The Tortugas region presents an ideal spot for studying the shallow-water
tropical Foraminifera of this particular region. It is removed from influence of shore
conditions; the water is at all times warm and pure, so that ecological conditions that are
present are constant. The twenty stations from which bottom samples were studied in the
preparation of this paper, together with collections from reef flats and from the eel-grass,
give a considerable range of conditions. The only stations at which Rotalia was found are
two in the moat at Fort Jefferson and the other in a very shallow lagoon at Long Key,
where the water was warm at low tide in June. By most authors, these specimens would
ordinarily be referred without question to Rotalia beccarii (Linnaeus). There are
differences from northern material, and in probability the Tortugas specimens belong to
different species. On the banks of dead coral which become exposed at spring tides, great
masses of attached Foraminifera develop. Of these, the most abundant is Homotrema,
which makes an appreciable contribution to the mass of material. With it, in crevices of the
dead coral, was a new species of Haliphysema. On the eel-grass (Posidonia), which forms
in shallow water inside the reef, there is Jridia, Planorbulina, Discorbis, Orbitolites, with a
peculiar miliolid which spreads over the surface. The mass of these must add appreciably
to the amount of carbonate of lime added to the bottom. The forms are rapid in their
growth, as the leaves of Posidonia are quickly covered in their growth by Foraminiferea
and other encrusting animals.
68. Cutright, P. E. 1937. Studies on the development of the dorsal spine of sting rays. Carnegie
Institution of Washington, Year Book 36: 90.
This report describes the collection of southern sting rays for a histological examination of
the stinging mechanism.
69. Dall, W. H. 1889. Reports on the results of dredging, under the supervision of Alexander Agassiz,
in the Gulf of Mexico 1877-78 and in the Caribbean Sea (1879-80), by the U.S. Coast
Survey Steamer "Blake":, Lieutenant Commander C.D. Sigsbee, U.S.N., and Commander
J.R. Bartlett, U.S.N. Commanding. XXIX- Report on the mollusca . Part II Gastropoda and
Scaphoda. Bulletin of the Museum of Comparative Zoology at Harvard College 18: 1-492,
with thirty one plates.
This listing of Mullusca collected by the “Blake” is supplemented by the southern
dredgings of the U.S. Fish Commission Steamer "Albatross" and other material collected
from the region. A systematic description and account of the gastropods and scaphopods is
given and illustrated. Nomenclature is discussed and rectified in several cases.
70. Darby, H. H. 1934. The mechanisms of asymmetry in the Alpheidae. Papers Tortugas Laboratory
28: 347-61 (issued Feb. 1934).
Carnegie Institution of Washington Publication Number 435.
In 1901, Przibram reported a series of striking experiments on the regeneration of chelae of
Alpheus dentipes, A. platyrhynchus and A. ruber. There is a pronounced quantitative and
qualitative difference between the right and left chelae. One chela is several times as large
as the other. Przibram showed that if the snap-claw is removed, at the next molt the pinch-
18
Hil
claw is changed into a snap-claw and a new pinch-claw is regenerated on the stump of the
old snap-claw. This unusual reversal of asymmetry was confirmed by Wilson (1903) and
Zeleny (1905). It was also shown that if both chelae are removed at the same time, they
regenerate in their original positions. These experiments seem to indicate that the final
degree of morphological expression of the gene may in certain cases depend on the
environment. In Crangon armillatus, an asymmetrical individual, it is symmetrical ten
days later. The morphological expressions of the gene are so concrete that it is difficult to
realize that the gene may also be the controlling agency in the production of definite
chemical substances, whose presence is manifested only by their physiological reactions.
Environment from that point of view can quite easily be thought to control the amount of a
substance produced. Changes might well be induced by radiation, due to the ionization of
the cell. Crangon has shown itself to be an organism in which studies on development as
an expression of the activity of the gene can be undertaken with some hope of success. The
nature of the regeneration of chelae in two members of the family of Alpheidae has been
studied; in particular, in Crangon armillatus. It has been shown that at certain stages in
the development of the chelae, a state is reached that permits the determination of which
side is to have the large chela, or snap-claw. Equal chelae have been produced
experimentally and are of three varieties: (1) both small (pinch-claws); (2) both large (snap-
claws); (3) both intermediate .
. 1940. Symmetry in normally asymmetrical crustacea. Papers Tortugas Laboratory 32: 61-
64 (issued Oct. 1939).
Carnegie Institution of Washington Publication Number 517.
A symmetrical specimen of Crangon armillatus was found in nature with two snap claws.
These claws differed in no way from snap claws produced experimentally and reported
previously.
72. Darby, H. H., E. R. F. Johnson and G. W. Barnes. 1937. Studies on the absorption and scattering of
solar radiation by the sea: spectrographic and photoelectric measurements. Papers
Tortugas Laboratory 31: 191-205 (issued Oct. 1936).
Carnegie Institution of Washington Publication Number 475.
A considerable amount of work has been done in recent times on the penetration of
radiation into sea and lake water. The importance of this work, in such matters as plant and
animal metabolism and under-water photography, is obvious. The amount and spectral
distribution of scattering, and the penetration of the ultraviolet component, are two phases
of the subject which have received scant attention. These studies were made from the
yacht, Elsie Fenimore at the Tortugas Laboratory. A comparative study has been made of
two methods of evaluating the transmission of various wave lengths of light through sea
water: (1) photometry by means of photoelectric cells, and (2) photographic
spectrophotometry. Bertel's observation that ultraviolet light penetrates a considerable
distance into the sea has been confirmed. The extent of penetration is greater than would
be expected from the laboratory data of Hulbert and Sawyer. A rough evaluation of the
transmissive exponent from 4500A to 3250A was made, which indicates the magnitude of
the disagreement. Scattering was found to be selective, becoming greater with decreasing
wave length. The spectral distribution of scattered radiation is indicated. The importance
of these observations for biological systems is outlined.
73. Davis, G. E. 1977. Anchor damage to a coral reef on the coast of Florida. Biological Conservation
11: 29-34.
Twenty percent of an extensive staghorn coral Acropora cervicornis has recently been
damaged by boat anchors in Fort Jefferson National Monument, Dry Tortugas, Florida. It
is suggested in this article that this type of damage may occur in other coral reef sanctuaries
74.
De
76.
19
unless anchor-sensitive areas are identified and closed to anchoring. Alternatively,
mooring buoys should be provided by sanctuary managers.
. 1982. A century of natural change in coral distribution at the Dry Tortugas, Florida USA. A
comparison of reef maps from 1881 and 1976. Bulletin of Marine Science 32, no. 2: 608-
73.
Changes in coral reef structure and composition at Dry Tortugas, Florida were compared
over a 95-year interval from benthic maps prepared in 1881 and 1976. Living hermatypic
corals occupied less than 4% of the 23,000-hectare area mapped, and showed little change
in area during the interval between maps. However, major changes in coral species
distributions and reef types were apparent. In 1976, a lush 220-hectare Acropora
cervicornis reef occupied what had been octocoral dominated hard bottom in 1881. The
44-hectare swath of A. palmata on the reef crest in 1881 was reduced to two small patches
totaling less than 600 m* in 1976. More than 90% of the extensive thickets of A.
cervicornis at Dry Tortugas were killed during the winter of 1976-77, apparently as a result
of thermal shock. These changes in coral distribution and abundance demonstrated the
natural dynamic nature of coral reefs, and showed the important role occasional short-term
extreme climatic events can play in shaping coral reef structure and species distribution.
The importance of protecting living corals and the value of ecosystem level sanctuaries as
dynamic standards are discussed.
. 1977. Effects of recreational harvest on a spiny lobster, Panulirus argus population.
Bulletin of Marine Science 27, no. 2: 223-36.
A commercially unfished population of Panulirius argus was studied in Fort Jefferson
National Monument at Dry Tortugas, Florida, from April 1971 to July 1975. For 29 months
all harvest was prohibited, then an experimental sport harvest (hand caught by recreational
divers) was allowed in 50% of the area for a period of 8 months, followed by 16 months of
complete protection for assessment of recovery. Data on the size, abundance, and natural
history of the lobsters were collected using SCUBA, and commercial trapping techniques.
A total of 4,257 lobsters, with a mean carapace length of 101 mm, was tagged and released
at Dry Tortugas. The existence of a resident adult P. argus population was demonstrated
by the recovery of all recaptured lobsters (7.3%) with 10 km of their respective capture
sites up to 104 weeks after release. Immediately following the experimental sport harvest,
the population in the sport harvested area showed a 58% reduction in trap catch rate and
dispersed to 42% of its pre-harvest lair occupancy density, while the population in the
unharvested control area remained essentially unchanged. The catch rate in the sport
harvested area recovered to 78% of its pre-harvest level after 1 year of complete protection
from harvest, and the lair occupancy rate recovery was 71% after 16 months of post harvest
protection. The pre-harvest standing crop was estimated at 58.3 kg/ha, wet weight.
. 1977. Fishery Harvest in an Underwater Park. Proceedings, Third International Coral Reef
Symposium, 605-8 no. 2. RSMAS, Univ. Miami, Coral Gables, Florida.
There is a potential conflict between park management for preservation of maximum
species richness and fishery harvest in parks. The recreational harvest of spiny lobster,
Panulirus argus, at Ft. Jefferson National Monument, Dry Tortugas, Florida, demonstrates
the nature and extent of the conflict. An eight-month-long diver harvest, limited by a daily
bag limit of two lobsters, reduced the previously unfished population by 585 and
significantly altered the local lobster distribution. Growth and natural recruitment did not
restore the population to its 58.3 kg/ha pre-harvest level, even after 16 months with no
additional harvest. The trophic status of spiny lobsters as high level carnivores and current
ecological theory combined with the harvest impact observed at Dry Tortugas suggests that
community structure and species richness would be significantly altered by the harvest.
20
77.
78.
19)
80.
81.
. 1975. Minimum size of mature spiny lobsters, Panulirus argus, at Dry Tortugas, Florida
USA. Transactions of the American Fisheries Society 104, no. 4: 675-76.
Of 1,594 female spiny lobsters examined during April 1973-1975 at the Dry Tortugas, 55%
were bearing eggs (berried). The specimens ranged in carapace length from 39 mm to 140
mm. No berried females were found with carapace lengths less than 78mm. Maturity was
reached by one half of the females in the 86-95 mm size class. The current minimum legal
size for sport and commercial lobster fishing in Florida is 76-mm carapace length.
. 1974. Notes on the status of spiny lobsters, Panulirus argus, at Dry Tortugas, Florida,
SUSF-SG-74-201. State University (Florida) System. Sea Grant Program. Publ..
Until mid-1971, sport harvest of spiny lobsters, primarily Panulirus argus, was permitted
in the 19,000 hectares underwater preserve created in 1935 which included the Dry
Tortugas atoll. At that time there was a two lobster per person per day limit. Few visitors
reached the isolated atoll during the first 20 years, with an average of some 1,200 people
per year. Annual visitation increased to over 21,000 in the late 1960's and early 1970's.
Concern was expressed for the protection of the quality and quantity of the lobsters found
in the area. The primary objective of the study was to assess the impact of human harvest
on a natural unperturbed lobster population.
. 1981. On the role of underwater parks and sanctuaries in the management of coastal
resources in the southeastern United States. Environmental Conservation 8, no. 1: 67-70.
Aquatic resources in parks and reserves are not as adequately protected as comparable
terrestrial resources. Thus the values of protected aquatic ecosystems as standards for
comparison, reservoirs of genetic materials, and ‘emotional’ reserves, are apt to be greatly
diminished. Even seemingly static ecosystems such as coral reefs are dynamic, changing
dramatically in response to natural short-term environmental variations. Such ecosystems
require protected natural areas as dynamic standards that will allow distinctions to be
drawn between effects of exploitation or pollution and normal variation. Furthermore,
fisheries harvests may reduce the size at which exploited species mature, and reduce the
amount and variability of genetic material produced by exploited populations. The seven
underwater parks or sanctuaries established since 1935 (Dry Tortugas) in Florida and the
U.S. Virgin Islands exhibit wide variations in the degree of protection accorded to aquatic
resources, a range being apparent from nearly complete protection in the first parks to be
established to virtually no protection at all in the recently established parks. The
consequences of permitting consumptive uses of aquatic resources in parks and reserves
need to be objectively evaluated. Unless these consumptive uses are severely curtailed or
eliminated, the primary values of the parks and reserves may never by realized.
. 1980. Spiny lobster series. Gary E. Davis (ed.), 27 pgs. American Fisheries Society:
Bethesda, MD.
This series of papers regarding spiny lobster management represents the efforts of a broad
cross section of the scientific fisheries community. Not only is there a diverse array of
disciplines from biochemical genetics to ecology and economics, but nearly every source
of research endeavor is represented. Members of two federal agencies (National Marine
Fisheries Service and National Park Service), a state agency (Florida Department of
Natural Resources), a public university (University of Florida), a private university (Nova
University), and a private company (Science Applications, Inc.) have combined their
efforts on a common subject that has already spawned thousands of scientific papers and
countless popular articles.
Davis, G. E. and J. W. Dodrill. 1989. Recreational fishery and population dynamics of spiny
lobsters, Panulirus argus, in Florida Bay, Everglades National Park, 1977-1980. Bulletin
of Marine Science 44, no. 1: 78-88.
21
Florida spiny lobsters Panulirus argus, occupied the southern two-thirds of Florida Bay in
Everglades National Park. Field studies of 3,570 tagged lobsters revealed that they pass
through Florida Bay, using it for less than three years as juveniles, between their planktonic
larval stages in the open ocean and adulthood on coral reefs. Lobsters from the bay
support commercial and recreational fisheries outside of Everglades National Park from
Dry Tortugas to Pacific Reef near Miami. Growth rates of juvenile lobsters in Florida Bay
are the highest on record, which may be a reflection of optimum habitat with abundant food
and shelter.
82. Davis, G. E. and J. W. Dodrill. 1980. Marine parks and sanctuaries for spiny lobster fisheries
management. Proceedings of the Gulf and Caribbean Fisheries Institution, pp.194-207,
32nd Annual Session.
National parks and sanctuaries with significant marine resources can play important roles
in effective fisheries management. However, if fishery resources are exploited and not
protected to the same extent terrestrial resources are protected in parks and sanctuaries,
they may not be available to provide the dynamic standards for comparison,
reproductive/genetic reserves, unique educational opportunities, and recreational escape.
Observations of more than 15,000 specimens of P. argus tagged at the National Park were
analyzed to provide data on migration patterns, natural mortality, reproduction and
development. Main factors affecting these populations were seasonality, stress in
juveniles, and sexual proportions in adults. Studies in non-exploited populations gave
good estimates of natural mortality. Size at first maturation was greater in non-exploited
populations than in exploited populations. Juveniles of P. argus show an extensive
directional migration pattern of 200 KM, while adults exhibit a restricted pattern for about
two years. Returned tags during the 1977-78 season in Florida came from sports fisherman
(49%), from commercial fisherman, (51%), and commercial traps (11%). This return
proves that sports catches were only 9% of the total in the northern part of the Florida Keys
(if all the tags were reported). The average of lobsters escaping from traps that were never
recovered was 1.2% daily, during the fourteen days that these were in operation.
83. Davis, J. H. Jr. 1940. The ecology and geologic role of mangroves in Florida. Papers Tortugas
Laboratory 32: 303-412 (issued Sept. 1940).
Carnegie Institution of Washington Publication Number 517.
The mangrove swamps of the low-lying coasts and islands of central and southern Florida
were studied during five seasons to determine the ecology of these unique littoral swamps,
and to obtain some idea of their importance as geologic agents in extending the coasts and
forming islands in the shoal-water regions. Five coastal and insular regions of the
peninsula were selected and a number of stations established in each for observations and
experimental studies. The report is divided into two parts. The "Ecology of the
Mangroves" is concerned with the types of plant communities of the mangroves and
associated vegetation, and the successional relationships of some of these communities.
"The Geologic Role of the Mangroves" considers the accretions of sedimentary and
cumulose soils in connection with the different agents that bring them about, and more
significantly, the role of the different mangrove communities in forming soils at higher and
higher levels. The most apparent succession of the mangrove communities consists of a
pioneer Rhizophora, a mature Rhizophora consocies, an Avicennia salt-marsh associes, not
always, flooded by salt or brackish water, a Conocarpus transition associes, seldom if ever
flooded by water, and a tropical or semitropical forest association, which is the actual
climax of the region. Besides Rhizophora, the Tortugas Keys have a young swamp of
Laguncularia around a pond on Bush Key, some young plants of Avicennia on Bush Key,
and a few old ones on Garden Key. Concocarpus was established on both Garden Key and
Bush Key. How these species got to the Tortugas and to many of the most isolated of the
Florida Keys is not certain, but should be considered.
727)
84. . 1942. The ecology of the vegetation and topography of the Sand Keys of Florida. Papers
Tortugas Laboratory 33: 113-95 .
Carnegie Institution of Washington Publication Number 524.
This is a study of the vegetation and some of the physiographic features of about thirty
islands of the Florida Keys, in an area extending west from Key West, Florida, and
including the Dry Tortugas Keys. About thirty islands of the Florida Keys beyond Key
West were investigated during the summer of 1940 and winter of 1942, and to some extent
during the summers of 1937 and 1938. These studies were concerned with the topography
and vegetation of these small, relatively isolated, and partly tropical islands. These islands
are here termed the Sand Keys because most of the parts above high tide are composed of
coarse calcareous sands, and also because this name was used by Willspaugh (1907). A
few of the Marquesas and Tortugas Keys have changed a great deal. The strand areas on a
number of islands seem to be increasing at the expense of the mangrove swamps. The
mangrove swamps have spread over wide areas in some instances and seem to be aiding in
building up the islands. Most of the constructional processes are, however, due to maritime
factors such as the ocean currents and tides. This paper is also a part of a series of studies
of the plant ecology of southern Florida. This and the author's study of mangrove
vegetation together describe most of the coastal and insular vegetation of that region.
85. Davis, R. A. Jr. and C. W. O'Neill. 1979. Morphodynamics of East Key, Dry Tortugas, Florida. in
Guide to Sedimentation for the Dry Tortugas, Fort Jefferson National Monument Florida
Southeast Geological Society Publication 21: 7-13.
East Key is comprised wholly of biogenic sand and fine gravel. It lacks beachrock or
bedrock which may act as a stabilizing agent such as on Loggerhead Key. During the past
two centuries, maps and charts documented the size, shape, and location of East Key. The
Key moved in a generally southeasterly direction across the shallow carbonate bank. East
Key was preserved, unlike some other islands, because of its easterly position with respect
to the deep lagoon. Those islands west of the lagoon moved easterly and disappeared into
the lagoon (O'Neill, 1976). East Key has decreased markedly in size during its
southeasterly movement. In addition there appears to be a change in morphology which is
related to seasonal changes in predominant wind direction.
86. de Laubenfels, M. W. 1936. A discussion of the sponge fauna of the Dry Tortugas in particular and
the West Indies in general, with material for a revision of the Families and Orders of the
Porifera. Papers Tortugas Laboratory 30: 1-225.
Carnegie Institution of Washington Publication Number 467.
Sponge specimens were collected near the Dry Tortugas by scientists affiliated with the
Carnegie Institution of Washington, or working at the laboratory maintained on
Loggerhead Key. These were sent to the U.S. National Museum to be studied by the
author. The West Indian region has long been known as one of the richest collecting
grounds for sponges in the world, and the Dry Tortugas offers a representative sample of it.
The author identified several new families and in many cases proposed new names for
families already in use. Representatives of each of the species discussed in this paper have
been deposited in the United States National Museum. Each new species is described in
detail.
87. ———. 1934. Physiology and morphology of Porifera exemplified by Jotrochota birotulata Higgin.
Papers Tortugas Laboratory 28: 37-66.
Carnegie Institution of Washington Publication Number 435.
The experimental work upon which this article is based was carried on during the summers
of 1927 and 1928 at the Tortugas Laboratory. A taxonomic description of the sponge was
provided. It was found that a hyaline ground mass or slime plays a very important role in
the life of Iotrochota and perhaps numerous other sponges. Judging only by items visible
23
in living Jotrochota cells, which were kept track of by conspicuously colored inclusions,
new sponges resulted from disassociated cells without intermediate differentiation and
respecialization. Reproductive bodies (gemmules) seem to result in Jotrochota by the
migration together of cells previously specialized for the purpose. Bispecific
conglomerations could be secured between Jotrochota and other species, and these
remained alive for two weeks or more, but whatever cell motility occurred within them
tended toward the ultimate segregation of the two species after somewhat the manner in
which animal gratings finally terminate. Amebocytes of Jotrochota sometimes ingested
flagellates which subsequently appeared as intracellular inclusions, and perhaps became the
symbionts whereby there occurred a certain amount of photosynthesis, the existence of
which was indicated by experimentation.
88.
. 1953. Sponges of the Gulf of Mexico. Bulletin of Marine Science of the Gulf and
Caribbean 2, no. 3: 511-77.
In 1948, a collection of sponges was made by the Marine Laboratory of the University of
Miami in the eastern Gulf of Mexico. Twenty-two stations were studied, at depths from 6
to 20 meters, in the area between Dry Tortugas and the northeastern part of the Gulf. The
collection comprises 52 species in 41 genera, all within the class Demospongea. Of these
11 species are new. Additional description is provided for a number of species. An
analysis of the sponge collection by stations is included.
89. de Renyi, G. S. 1934. Studies of nerve cells of invertebrates. Carnegie Institution of Washington,
Year Book. 33: 250.
The nerve tissue (neuroplasm) of gastropods (Strombus gigas), Aplysia protea, Olivia
litterata, Cypraea exanthema, Casio cameo), decapods (Panulirus argus, Crangon
armillatus, Ocypoda albicans), and hemichordates (Ptychodera bahamensis) were studied.
The neoplasm of the Gastropoda and Hemichordata exhibited viscosity, and a certain
degree of elasticity, whereas decapodean neuroplasm was liquid.
90. Deflaun, M. F. 1987. "The distribution and molecular characterization of dissolved DNA in aquatic
environments." University of South Florida. Ph.D. Dissertation
The distribution of dissolved DNA in oceanic, estuarine and freshwater environments in
southwest Florida and the Gulf of Mexico was determined by using a method for the
measurement of dissolved DNA based on the fluorescence of Hoechst 33258-DNA
complexes. Oceanic concentrations of extracellular DNA ranged from 0.2 to 19
decreasing as a function of distance from the shore and depth in the water column.
Samples of the mucus-rich coral surface microlayer (CSM) collected on reefs in the Dry
Tortugas had dissolved DNA concentrations from 1.8 to 11.7 times that in the overlying
water. Estuarine concentrations, measured at three stations in Tampa Bay, FL over a 15-
month period, followed the seasonal trend in concentrations in offshore environments,
while variations in the estuary were significant, with maximum concentrations in nighttime
samples. Although concentrations of dissolved DNA in the eutrophic Alafia River were
generally higher than those in the oligotrophic Crystal River, values as low as 1.14 were
measured in the Alafia. A wide range of molecular weights (determined by agarose gel
electrophoresis) was found for extracellular DNA concentrated from various aquatic
environments. These results indicated that dissolved DNA is in a size range sufficient to
contain gene sequences, which may be important in natural transformation of microbial
populations. A model system for probing extracellular DNA from aquatic environments
was developed using the plasmid containing the herpes simplex thymidine kinase (TK)
gene. Plasmid DNA and the TK gene fragment added to artificial seawater were
concentrated and labeled TK to establish percent recovery and detection limits for the
method. The degradation of plasmid DNA added to a natural seawater sample was
monitored over a 36 h period by probing with the TK gene probe. Intact plasmid was
24
detected for up to 4 h and DNA hybridizable to the TK probe was detected for up to 24 h.
These methods were used to probe for the TK gene in environmental samples of
extracellular DNA. Hybridization to the TK probe was detected in both freshwater and
estuarine samples.
91. Dinsmore, J. J. 1972. Sooty tern behavior. Bulletin of the Florida State Museum of Biological
Science 16, no. 3: 129-79.
A four-year study of the breeding behavior of sooty terns (Sterna fuscata) was made at
Bush Key, Dry Tortugas in the southeastern Gulf of Mexico. The results are compared
with the behavior of other terns and the differences discussed, particularly in regard to the
pelagic environment the sooty tern inhabits. sooty terns have a lower clutch size, longer
period of development of the chick, and first breed when older than most other terns, many
of which feed in marshes and coastal waters. These characteristics of sooty tern breeding
biology are similar to those of many other pelagic birds. A distant food supply and high
adult survivorship apparently have contributed to these differences from other terns.
92. Dinsmore, J. J. and W. B. Robertson Jr. 1972. Sooty tern feeding on moths. The Auk 89, no. 2:
93. Dole, R.
440.
While banding sooty terns (Sterna fuscata) at Bush Key, Dry Tortugas, Florida on June 28,
1970, an adult tern regurgitated two moths 1.5 to 2 cm long together with several
unidentified fish. The moths were identified to the family Noctuidae. Although the food of
sooty terns at the Dry Tortugas has not been studied in detail, sizable collections of food
regurgitated show that this population feeds on fish and squid. In 13 years of tern banding,
this is the first time an insect has been found as part of the sooty tern's diet .
B. 1914. Some chemical characteristics of sea water at Tortugas and around Biscayne Bay,
Florida. Papers Tortugas Laboratory 5: 69-78.
Carnegie Institution of Washington Publication Number 182.
The chemical tests at Tortugas were performed by the writer in June 1913, in the Marine
Biological Laboratory, Tortugas, Florida, for the primary purpose of ascertaining what
soluble effect, if any, carbon dioxide in sea-water might have on coral and other deposits of
calcium carbonate. The tests of waters from Biscayne Bay were made to ascertain the
differences in concentration of sea-water in the bay and the diluting effect of Miami River.
The salinities of the three samples taken outside the reefs agree closely with each other and
with the salinity of Gulf water at Tortugas, Florida (36.01 ppt), which is somewhat greater
than that of standard ocean water (35.02 ppt.). The water in the south part of the bay is
somewhat more concentrated having salinities of 36.73, 36.64, and 36.64 ppt., respectively.
This evidence that the water in this part of the bay is concentrated by evaporation during its
retention in the shallows serves further to indicate that circulation there is not very rapid
and that the greater bulk of the water inside the keys is not thoroughly mixed or shifted by
the tides. Sample 1 has a salinity obviously higher than the pure water of Miami River
alone may be expected to have, and represents admixture with bay water; carbonates are
absent from it, but bicarbonates are much higher than in the normal drainage from the
Everglades and may be attributed to reaction of the carbon dioxide that the river water
carries.
94. Domeier, M. L. Speciation in the Serranid fish Hypoplectrus. Bulletin of Marine Science 54, no. 1:
103-41.
Research was conducted to determine the species status of individual color morphs of
fishes in the genus Hypoplectrus (family Serranidae). Crossing two morphs of
Hypoplectrus (H. unicolor x H. gema) in the laboratory produced an F1 generation with an
intermediate phenotype to that of the parental types. This intermediate morph cannot be
assigned to any known morph and is thus termed a hybrid. Individuals of several
25
Hypoplectrus morphs were found to select only individuals of the same morph as a mate
when provided a choice. Individual fish can sometimes be forced to mate with an
individual of a different morph by not providing a choice of mates. The occurrence of
hybrids was found to be low in the field, corresponding to the low occurrence of mixed
matings in the field. Some differences in distribution were found between the different
hamlet morphs. The new data provided by this study, which includes specimens collected
from the Dry Tortugas, indicate that the different color morphs warrant full species rank. It
is hypothesized that speciation in Hypoplecturs was driven by the rise and fall of sea level
during the last ice age.
95. Donaldson, H. H. 1916. Experiment on the feralization of the albino rat. Carnegie Institution of
Washington, Year Book 15: 200-201.
Domesticated albino Norway rats were released on East Key to determine if changes in
brain weight occur over successive generations in a wild state. Since the rats were
unmarked, it was impossible to ascertain if differences in weight were from new breeds or
from animals in the original colony.
96. Doyle, W. L. 1936. Cytology of Valonia. Papers Tortugas Laboratory 29: 13-21 (issued Nov.
SHE
1935).
Carnegie Institution of Washington Publication Number 452.
For a number of years algal cells with large vacuoles have been the subject of research on
the permeability of the plasma membrane. Prominent among forms investigated is
Valonia. This paper describes the cytology of Valonia ventricosa and Valonia macrophysa
with particular emphasis on structures of significance in physiological investigations. The
cells were collected on Bush Key Reef and from the moat at Fort Jefferson and kept in
finger bowls in the laboratory. The morphology of the various structures in the cytoplasm
of Valonia macrophysa and V. ventricosa is described. The plastids produce starch and
lipoid granules and are sufficiently numerous as to constitute two-thirds-of the volume of
the cytoplasm. There are approximately three hundred nuclei per square millimeter of cell
surface in the coenocytes. Mitosis is intranuclear. In the development of the rhizoidal
hapteron cells of the aplanospores, the mitochondria arise from plastids of the coenocyte in
which the aplanospores were formed. The large central vacuoles of the coenocyte arises by
fusion of small vacuoles formed in the cytoplasm. Double vital staining of artifact
vacuoles is noted.
. 1940. The structure and composition of Valonia ventricosa. Papers Tortugas Laboratory
32: 143-52 (issued Sept. 1940).
Carnegie Institution of Washington Publication Number 571.
The physiology of Valonia has been dealt with extensively by numerous authors. Cells
were collected from Long Key and the adjacent reef and brought to the laboratory, where
they were kept in large glass jars of sea water which was changed daily. Measurements
have been made of the relation of the volume and thickness of the cytoplasm and cell wall
to the size of the coenocyte. The specific gravities of various cell constituents and of cells
of various sizes have been measured. From a consideration of the results presented it
would appear that the level of metabolic rate in Valonia is of a low order, but not
necessarily of a different order of magnitude from that of the barley-root and potato-slice
systems.
98. Doyle, W.L. and M. Metcalfe Doyle. 1940. The structure of zooxanthellae. Papers Tortugas
Laboratory 32: 127-42 (issued May 1940).
Carnegie Institution of Washington Publication Number 517.
The structure of the zooxanthellae in various invertebrate reef organisms under various
conditions was investigated at the Dry Tortugas in 1934, 1935 and rechecked in 1939. Ten
26
species of corals and foraminifera were studied in their living conditions, as well as after
fixation. Zooxanthellae in foraminiferans, were examined for the effects of light in normal
gas tensions, in increased carbon dioxide tensions, and on specimens in oxygen and
hydrogen; while in corals the comparative cytology of zooxanthellae was studied. For the
large heads of the Orbicella (Madrepora), the amount of light present at the top and
bottom of the corals determined the natural variations in the amount of calcium oxalate
crystals in zooxanthellae. Increased levels of crystals were found at the bottom in
darkness, while no crystals were found at the top. Similar results were found in
foraminiferans. The converse is true for the amount of starch present. The zooxanthellae
in corals under the most intense natural light conditions contains little starch, but abundant
oil droplets. It was concluded that, overall, for the greater part of the day, the
zooxanthellae, as well as the corals, are in need of oxygen.
99. Drew, G. H.. 1914. On the precipitation of calcium carbonate in the sea by marine bacteria, and on
the action of denitrifying bacteria in tropical and temperate seas. Papers Tortugas
Laboratory 5: 7-45.
Carnegie Institution of Washington Publication Number 182.
The investigations described in this paper were made in the summers of 1911 and 1912
under the auspices of the Carnegie Institution of Washington. The intent was to study the
action of marine denitrifying bacteria in tropical seas. The discovery that these denitrifying
bacteria also possess the power of precipitating calcium carbonate from soluble calcium
salts present in sea-water has overshadowed the primary object of the work. The
observations so far available are few, and the area they cover too small, to attempt to make
broad generalizations. However, it can be stated that the very extensive chalky mud flats
forming in the neighborhood of the Florida Keys are now being precipitated by the action
of the bacterium calcis on the calcium salts present in solution in sea-water. The
investigation can at most be considered to offer a mere indication of the part played by
bacterial growth in the metabolism of the sea. To obtain a real insight into the question, it
would be necessary to make more extensive bacterial and chemical observations in
tropical, temperate, and arctic waters, to study the bacteriology of other areas where
calcium carbonate is being precipitated from the sea, and to make further investigations in
the laboratory into the chemistry of the reactions that can be brought about by various
species of marine bacteria.
100. Dustan, P. 1985. Community structure of reef-building corals in the Florida Keys , Carysfort Reef,
Key Largo, and Long Key Reef, Dry Tortugas. Atoll Research Bulletin 282-292: 1-29.
This communication is the result of two parallel studies on the distribution of reef-building
corals on Carysfort Reef, Key Largo and Long Key Reef, Dry Tortugas. The aim of the
projects was to characterize the species composition of reef-building corals from the
northern and southernmost localities of the Keys, and through comparison attempt to
identify the impact of man on the reefs in the Key Largo area of the northern Florida Keys.
101. Dustan, P., W. Jaap and J. Halas. 1976. The distribution of members of the Class Sclerospongiae.
Lethaia 9, no. 4: 419-20.
The Sclerospongiae play an important and sometimes major role in the construction and
infilling of reefs in tropical waters. Modern sclerosponges are limited to dark, quiet,
sediment-shaded habitats. This study describes the distributions of sclerosponges in the
Bahamas and the Florida Reef Tract. The sponges were found in the Grand Bahamas.
After extensive SCUBA diving in Pennekamp Park and the Dry Tortugas, no
Sclerospongiae were found. Cold water temperature, or alternatively few, if any larvae to
colonize the reef tract are possible explanations for the lack of Sclerospongiae in the
Florida Reef Tract.
27
102. Edmondson, C. H. 1908. A variety of Anisonema vitrea. Papers Tortugas Laboratory 1: 191.
Carnegie Institution of Washington Publication Number 102.
Notes are provided on the protozoan, Anisonema. Anisonema vitrea (Dujardin) is a
flagellated protozoan, elongate-oval in form, the anterior end broadly rounded, the
posterior more acutely rounded. Anisonema vitrea is distinguished from other species of
the genus by eight furrowed surfaces extending in a slightly spiral manner from one end of
the body to the other. During the summer of 1906, while working on marine Protozoa at
the Tortugas, Fla., the author studied a form considered as a variety of the above species
entitled Anisonema vitrea (Duj.) var. pentagona. A description of the difference between
the species and variety is presented .
103. Erseus, C. and M. R. Milligan. 1988. A new Bathydrillus oligochaeta Tubificidae from the eastern
Gulf of Mexico. Bulletin of Marine Science 42, no. 2: 292-95.
Bathydrilus natabilis is described from 4-58.5 meter depths off Crystal River and Dry
Tortugas in the eastern Gulf of Mexico. The species is characterized by large, finely
pectinate, penial setae in segment 11 and entally curved, single-pointed, spermathecal setae
in segment 10 which distinguish it from all congeners.
104. Farfante, I. P. 1980. A new species of rock shrimp of the Genus Sicyonia penaeoidea, with a key to
the western Atlantic species. Proceedings of the Biological Society of Washington 93, no.
3: 771-80.
Sicyonia olgae, new species, ranges from Dry Tortugas Islands, Florida, to Surinam. It
differs from Sicyonia typica (Boeck, 1864), its closest western Atlantic relative, in
possessing sublateral carinae on the carapace, and in lacking posterior pleural sulci on the
first three abdominal somites. Also distinctive are the sharply pointed, mesially directed,
distomesial projections of the petasma in the male, and in the female the pair of long,
slender spines on sternite XI and rounded posterolateral processes of the median plate of
sternite XIII. A key to the western Atlantic species of Sicyonia is supplemented by
synopses of their geographic and depth ranges which include many extensions of
previously known limits.
105. Feinstein, A. A. A. R. Ceurvels R. F. Hutton and E. Snoek. 1955. "Red tide outbreaks off the
Florida West Coast." Report to the Florida State Board of Conservation of Marine
Laboratories .
A compilation of reports of red tide on the west coast of Florida from 1844 to January,
1955 is given. Also included are two working diagrams of incidence of red tide,
suggesting that red tide occurs more frequently in the months of August through January,
and that individual red tide outbreaks are part of larger outbreaks, which seem to move
from south to north, and summer outbreaks appear to originate mostly north of Venice,
winter and spring outbreaks further south. Further data are required to give complete
support. If this is substantiated, control may be exerted by action in a limited focal area or
areas of origin. Otherwise, the problem of control may be of the greatest difficulty, since it
will require action over a much wider area.
| 106. Field, R. M. 1919. Investigations regarding the calcium carbonate oozes at Tortugas, and the beach
rock at Loggerhead Key. Carnegie Institution of Washington, Year Book 18: 197-98.
Calcium carbonate accumulations in the shallow lagoons and channels between the reef
flats were examined to ascertain their origin. Carbonate ooze hardens rapidly when
exposed to air and when flooded with saltwater, mud-cracked zones can be formed similar
to those in the geologic record, as in the Stones River limestone formation. An account is
given on the origin of the "beach rock" found between the high and low water marks on
Loggerhead Key.
28
107. :
1920. Origin of "beach rock" (coquina) at Loggerhead Key, Tortugas (abs.). Bulletin of the
Geological Society of America 31: 215.
A study was made to discover the origin of the "beach-rock" or cemented shell-sands
which occur between high and low tides. By means of a stand-pipe and pump, it was found
that during heavy rains a shell key acts like a reservoir, and the meteoric water dissolves
CaCO; on its way through the loose shell sands. The ground water was found to contain 40
per cent more CaCo; in solution, or colloidal suspension, than the normal sea water. This
concentrated solution of CaCo; has a strong cementing value, and is probably an important
factor in the formation of the "beach-rock" where the ground water flows out through the
beach sands, between tides.
108. Fisk, E. J. 1976. Black phoebe sighted at Dry Tortugas. Florida Field Naturalist 4, no. 2: 39.
An observation of a black phoebe on Loggerhead Key, Dry Tortugas on April 13, 1976 is
recorded. This is the fourth sighting and only spring record for Florida of a black phoebe.
109. Gauld, G. 1820. An accurate chart of the Tortugas and Florida Keys or Martyrs, surveyed by
110. Gee, H..
George Gauld, A.M. in the years 1773, 1774, 1775. London, W. Faden.
First nautical chart of the Dry Tortugas is produced.
1934. Lime deposits and the bacteria. I. Estimate of bacterial activity at the Florida Keys.
Papers Tortugas Laboratory 28: 67-82 (issued Dec. 1932).
Carnegie Institution of Washington Publication Number 435.
Aerobic organisms were collected from the Florida Keys. Viable counts indicate that open
areas are only thinly populated with these forms, but that sheltered areas may permit
increased activity. Conditions in the mud are such as to favor the growth of anaerobes.
There is a possibility that specific groups, such as the purple sulphur organisms are at work
in addition to the conventional aerobes.
111. Gee, H. and C. B. Feltham. 1934. Lime deposition and the bacteria. II. Characteristics of aerobic
112. Gersh, I.
113. Gilmore,
bacteria from the Florida Keys. Papers Tortugas Laboratory 28: 83-91 (issued Dec. 1932).
Carnegie Institution of Washington Publication Number 435.
General bacterial conditions at the Florida Keys during the 1930 season have been
discussed by Gee (1932). There was reported a collection of 138 representative aerobic
organisms recovered from the water and mud of Bird Key harbor between Bird and Garden
Keys, of the Marquesas lagoon, and of one vertical one in the vicinity of the Gulf Stream.
Preliminary examinations were made of them at the Tortugas laboratory. The strains were
found to be Gram-negative rods, ammonia-producing, and possibly fermenting. The
collection was subsequently studied exhaustively at the Scripps Institution during the
winter of 1930-31. When freshly isolated, these bacteria displayed considerable variation
in size, in colony features and color, and in their degree of physiological activity.
1935. Studies on the anterior pituitary gland of the nurse shark. Carnegie Institution of
Washington, Year Book 34: 81.
Experiments were planned on the nurse shark to determine which of the activities of the
anterior pituitary gland are referable to the eosinophile cells.
R. G. and R. S. Jones. 1988. Lipogramma flavescens, a new grammid fish from the
Bahama Islands with descriptive and distributional notes on L. evides and L. anabantoides.
Bulletin of Marine Science 42, no. 3: 435-45.
In 1981, dredge collections made north of the Dry Tortugas by Continental Shelf
Associates under contract with the Bureau of Land Mangement documented the first
continental record of L. anabantoides.
29
114. Ginsburg, R. N. 1953. Beach rock in south Florida. Journal of Sedimentary Petrology 23: 89-92.
115. Goldfarb
116.
IIY/,
The rapid intertidal lithification of beach deposits in the coral seas has received the
attention of numerous investigators. Study of beach rock from the Dry Tortugas shows that
the aragonite cement is precipitated from the sea water remaining in the beach sands at low
tide. High temperatures, rate of beach drainage, and the permanence of the beach control
the localization of beachrock. The recognition of beachrock in the fossil record is briefly
discussed.
, A. J. 1913. Changes in concentration of sea-water and their influence upon regeneration.
Proceedings of the Society for Experimental Biology and Medicine 10, no. 3.
The regeneration under changed densities of sea water was observed under conditions that
endured the elimination of uniformity of associated factors such as size of medusae,
volume, surface and depth of solutions, extent of injury, level of amputation, temperature,
crowding, aeration, etc. Dilutions were made with water containing a known quantity of sea
salts, and concentrated solutions were made by slow evaporation, which corrected certain
errors in previous experiments. Results were compared with those of Loeb. It was found
that both the hydroid Eudendrium of Woods Hole as well as Cassiopeia of Dry Tortugas
differed radically from the Loeb experiments, in respect to the range of solutions in which
animals lived or regenerated, the optimum solutions, the normality of the regenerated parts
and the character of the curve. It is stated that Loeb's curve probably is limited to
Tubularia of Naples, and does not represent the behavior of organisms to changes of
density of sea water, and that the differences in the behavior of these three organisms can
hardly be correlated with the differences in concentration of the sea water in which they
normally live.
. 1914. Changes in salinity and their effects upon the regeneration of Cassiopea xamachana.
Papers Tortugas Laboratory 6: 83-94.
Carnegie Institution of Washington Publication Number 183.
Cassiopea xamachana, a large scyphomedusa, is very abundant in the very shallow waters
of the moat at Fort Jefferson, Dry Tortugas, Florida. The present report considers to what
extent changes in salinity influence regeneration in Cassiopea, and the results of the
investigation are compared with those previously obtained with the hydroid Eudendrium
ramosum of Woods Hole, Massachusetts and with the observations of Loeb with the
hydroid Tubularia of Serino Bay, Italy. The object of this investigation was to ascertain to
what extent changes in salinity affected Cassiopea xamachana normally subject to
relatively great variation in the concentration of the sea-water, and to compare the results
with those of the hydroid Eudendrium and the hydroid Tubularia. The following variable
factors were uniform for the series: size of medusae; volume, surface, and depth of the
solutions; extent of injury; level of amputation; temperature; crowding. Injurious or other
variable factors were guarded against. Cassiopea lived in solutions ranging from 40 to 153
per cent sea-water solutions. Regeneration occurred in solutions containing 50 to 133 per
cent sea-water. Normal regeneration of the arms occurred within much narrower range,
namely 75-105 per cent. Beyond these limits regeneration was atypic .
. 1918. Effects of aging upon germ cells and upon early development. Part II. Biological
Bulletin 34, no. 6: 372-409.
In a previous preliminary experiment it was shown that freshly liberated eggs of different
females of three different species of sea urchins (Toxopneustes and Hipponoe collected
from the Tortugas, and Arbacia from Woods Hole, Mass.) varied in respect to size, jelly
layer, membrane formation, and cleavage. In this paper the same technique and the same
three species of sea urchins were used to determine the physiologic condition of the germ
cells, and then determine the nature of the changes in the eggs as they became increasingly
overripe. As eggs in good physiologic condition aged, their volume increased until they
30
118.
120.
121.
became smaller than the norm. Eggs in poor condition were reduced in size., in all three
species, there was a loss in jelly layer with age, depending on the condition of the egg. In
all three species, as the eggs aged, the membrane appeared closer to the surface, becoming
thinner until none was formed. The rate of decrease in cleavage with age was greater in
Toxopneustes and Hipponoe than in Arbacia. Overall, the change in size, jelly membrane,
and cleavage with aging of germ cells are accurate, convenient and corroborative indices
of physio-chemical and morphologic changes in the eggs as they age, and afford convenient
measures of loss in vitality, or physical deterioration.
. 1914. Experimentally fused larvae of echinoderms with special reference to their skeletons.
Papers Tortugas Laboratory 6: 103-21.
Carnegie Institution of Washington Publication Number 183.
The early work of Loeb, Morgan, and Herbst on the production of multiple embryos from a
single egg suggested the reverse experiment of grafting or reuniting several fertilized eggs
into on embryo. In 1912, the writer repeated these experiments with the American form
Arbacia punatulata and succeeded only after slightly modifying Driesch's method.
Subsequently, in the performance of other experiments, it was discovered that eggs could
be agglutinated and fused quite as readily by a very different method, which was not only
simpler but free of certain objections that might be urged against previously known
methods. The new method consisted in using an isotonic or slightly hypotonic NaCl
solution diluted with varying quantities of sea-water.
. 1913. The influence of the central nervous system in regeneration of an annelid worm.
Proceedings of the Society for Experimental Biology and Medicine 10, no. 3. np.
(No abstract available).
. 1914. Regeneration in the annelid worm Amphinsoma pacifica, after removal of the central
nervous system. Papers Tortugas Laboratory 6: 95-102.
Carnegie Institution of Washington Publication Number 183.
In a previous publication, the writer found that the head of the earth-worm Lumbricus was
regenerated in the entire and permanent absence of the nerve-cord from the amputated
region. The marine annelid worm Amphinoma pacifica readily regenerated a head at all
levels except the distal eighth of the worm. Regeneration may be prevented by a severe
injury, either to the digestive tract or to the central nerve system; the greater the injury the
more likely will regeneration be inhibited. Many pieces did not regenerate after removing
the alimentary tract from five or more segments nearest the amputated level. Many pieces,
about one-third, failed to regenerate after removing the nerve-cord by the forceps, i.e., with
little injury to adjoining tissues. All failed to regenerate after removing the nerve-cord by
the "window" method. The operated worms were examined in serial sections. In one group
a regenerated nerve-cord connected the regenerated "brain" and commissures with the old
intact nerve-cord. In a second group the regenerated nerve-cord approached and in
instances reached the amputated level, yet no head was formed. In a third group, the nerve-
cord had regenerated, but several segments nearest the amputated end were yet without any
nerve-cord or ganglia. These worms nevertheless had regenerated a head with its typical
brain and nerve-commissures.
. 1917. Variability of eggs and sperm of sea-urchins. Papers Tortugas Laboratory 11: 71-87.
Carnegie Institution of Washington Publication Number 251.
A clear understanding of the variability in normal fresh eggs and sperm is necessary in
order to appreciate and to evaluate the changes that take place in overripe germ-cells. This
paper deals exclusively with the qualitative and quantitative differences of such freshly
collected sea-urchin eggs and sperm and with the differences in their early development.
Sil
122. . 1917. Variability of germ cells of sea urchins. Proceedings of the National Academy of
Science 3: 241-45.
Three different species of sea urchins (Toxopneustes and Hipponoe collected at the Dry
Tortugas, Arbacia collected at Woods Hole, Massachusetts) were used to determine the
normal variability of sea urchin germ cells. Having determined the optimum and constant
conditions of germ cells, studies were conducted to examine variations in size and shape of
eggs, the jelly layer of eggs, membrane formation, and cleavage among the three species.
Amazingly large variations were found in fresh germ cells among species, thus suggesting
that among other investigators of the varying behavior of the eggs, a large part of the
variation was probably due to the physiologic conditions of the eggs which these
investigators used.
123. Goodrich, H. B. 1935. Color patterns in fish. Carnegie Institution of Washington, Year Book 34: 81.
Studies were carried out to investigate internal conditions which may control the
development and maintenance of color patterns in fish by transplanting scales and tissues
from one type of pigment area to another.
124. Gordon, M. 1933. The internal pigment systems of fishes. Carnegie Institution of Washington, Year
Book 32: 268.
The internal pigmentary systems of major taxonomic groups were examined. Halichores
bivittatus and Lutjanus griseus showing possibly neoplastic growths were collected for
study.
125. Goy, J. W. 1982. West Indian Stenopodidae. 2. Occurrence of Richardina spinicincta Crustacea,
Decapoda, Stenopodidea off the Dry Tortugas. Bulletin of Marine Science 32, no. 1: 344-
47.
An examination of Richardina spinicincta collected by W.L.Schmitt in August of 1932 is
made. It is concluded that this specimen is truly R. spinicincta, that this is the sixth known
specimen of the species, and the first record of the genus in the Western Atlantic. The
occurrence suggests that the genus occurs at shallower depths than those recorded in
previous literature.
126. Grave, C. 1934. The Botryllus Type of Ascidian larva. Papers Tortugas Laboratory 28: 143-56
(issued Dec. 1932).
Carnegie Institution of Washington Publication Number 435.
Free-swimming larvae of at least three well-defined types are found in life cycles of
ascidians; one, characteristic of species of Molgula (Grave '26) and related genera, that
has one sense organ only, a statolith, in its sensory vesicle. The nerve cord lies in a mid-
dorsal position above the notochord, the caudal fin is expanded vertically in the median
plane and adhesive papillae are lacking. In the text the structural organization of a type of
larva characteristic of species of Botryllus and related genera is described. The body is egg
shaped, its depth being approximately the same as its width. Three conical sensory
papillae arranged in the form of an equilateral triangle are borne at the anterior smaller end
of the body, two located on either side of the median plane dorsal to the central body axis,
one in the median plane ventral to the central axis. The same gross parts found in the
Central nervous system of larvae of other types are present. The anterior end of the visceral
ganglion bends to the right and expands to form the sensory vesicle, which, in contrast with
that of larvae of other types, does not project to the level of the dorsal surface of the body
but retains an interior position relatively far below the surface.
127. ———. 1936. Metamorphosis of ascidian larvae. Papers Tortugas Laboratory. 29: 209-91 (issued
Dec. 1935).
Carnegie Institution of Washington Publication Number 452.
32
The studies of metamorphosis of larvae of ascidians were made during the summers of
1927, 1930, and 1933 at the Tortugas Laboratory with the purpose of finding methods of
accelerating and controlling metamorphosis and thus of discovering something of the
fundamental nature of the internal mechanism involved and the environmental conditions
with which it is causally related. The observations made in the cqurse of this investigation
are interpreted as follows: The ascidian larva is a dual organism, the action system of the
larva being quite separate from the action system of the ascidiozooid. Metamorphosis
advances by three stages; (a) changes in the adaptive responses of the larva to light and
gravity; (b) the attachment of the larva to the surface of some foreign object; (c) the
disruptive phase during which the entire larval action system is destroyed. Swimming
activity causes the production and concentration of some metabolic product in the larval
tissues that is essential to the induction of metamorphosis. The presence in the larval
tissues of metabolic products resulting from swimming is not alone sufficient to induce
metamorphosis, but another substance with which this metabolic product may react is
equally necessary. The great variability of ascidian larvae of the same species in the
duration of their free-swimming period is apparently due to variability in the time of
formation of the susceptibility substance and hence to the time of differentiation of the
larval organ that produces it. Metamorphosis may be induced artificially by diverse
chemical and biological substances placed in sea-water with groups of larvae in lactic acid.
Metamorphosis is rapidly and consistently induced in the larva of Phallusia nigra. The
activating agents extracted from the fresh ascidian tissues that were so specific in their
effects may also be endosymes of a highly specialized kind, each found only in a single
species of ascidian. The mechanism of metamorphosis is comparable in its organization to
that of development of an egg, which also may be activated by numerous and diverse
chemical and physical agencies.
128. Grave, C. and P. A. Nicoll. 1940. Studies of larval life and metamorphosis in Ascidia nigra and
species of Polyandrocarpa. Papers Tortugas Laboratory 32: 1-46 (issued Oct. 1939).
Carnegie Institution of Washington Publication Number 517.
Experimental studies made during the summer of 1933 (Grave, 1936) demonstrate that sea-
water extract of pharyngeal, atrial, or mantle tissues of adult Ascidia nigra is effective in
inducing 100 per cent metamorphosis in groups of Ascidia larvae within 3 hours after
hatching and that similar extract of tissues of Polyandrocarpa induces 100 per cent
metamorphosis in groups of Polyandrocarpa \arvae within 42 minutes after liberation
from the parent colony. These observations led to a search during the summers of 1935
and 1936 for a specific chemical substance in the tissues of these ascidians having the
properties required for the rapid acceleration of the process of metamorphosis. An account
of the methods and results of this work is given in this paper. It was found that the amino
acids |-histidine, leucine, glycine, cysteine, and d,]-alanine, in the form received from the
laboratories in which they were prepared, accelerated metamorphosis in groups of larvae of
both types. A sea-water extract of free-swimming larvae or of late embryonic stages of
Ascidia has the same accelerating effect on metamorphosis of Ascidia larvae as an extract
made from tissues of the adult ascidian. Heating adult Ascidia tissue or releasing distilled-
water extracts of the tissue for several hours does not destroy the accelerating substance.
Non-toxic concentrations of copper, iron, and aluminum salts induce early metamorphosis
to a marked degree. The duration of the free-swimming period of Ascidia larvae is longest
at the beginning of the breeding season of the species and becomes gradually shorter as the
season advances.
129. Gudger, E. W. 1921. Notes on the morphology and habits of the nurse shark, Ginglymostroma
cirratum. Copeia 98: 57-59.
A physical description of the nurse shark as observed by the author for several summers in
the southern Florida Keys and Dry Tortugas is given.
130.
Wil,
N32,
133),
33
. 1929. On the morphology, coloration, and behavior of seventy teleostean fishes of
Tortugas, Florida. Papers Tortugas Laboratory 26: 149-204.
Carnegie Institution of Washington Publication Number 391.
In the course of work at the Tortugas, 70 teleosts, belonging to 28 families, have been
studied. Habits have been recorded herein that stand out prominently to the taxonomist.
First, basing the classification of tropical fishes on coloration is a very dangerous thing.
Most of the fishes in Tortugas have from two to five color phases in life and, even when
studying the fish in a state of comparative quiet in an aquarium, it is very difficult to
determine which is its normal color. When a fish dies, its color changes either entirely or in
its intensity, so that the coloration of the dead fish is markedly different from that of the
live fish. It is equally dangerous to describe and classify a tropical fish from a single
specimen, since these fishes are so very variable in the number of fin rays, in the relative
proportions of the body, in scale count, and in the many details which help to distinguish
one species from another..
. 1918. On the use of the diving helmet in submarine biological work. American Museum
Journal 18: 135-38.
The use of the diving helmet for research at the Dry Tortugas was initiated in 1915 by
Longley and Carey, for fish observations and photography. Its use was declared new for
underwater work. However, such is not the case. The use of the helmet alone replaced
cumbersome diving suits (scaphanders) used by the commercial spongers out of Tarpon
Springs, Florida, and early workers on the construction of the overseas railroad, The
Florida East Coast Railway Extension from Homestead to Key West, Florida. The diving
helmet in biological work dates back to around 1845, when M. Milne-Edwards conducted
bottom surveys off the coast of Sicily. In 1679, pressurized air was first supplied to
Borelli, who attached a simple air compressing pump to a leather diving helmet. These
devices are all refinements of the crude diving helmets used back in ancient times by
Alexander the Great, while recording plant and animal observations. These are some of the
earliest underwater biological observations ever recorded. The earliest account of any type
of diving apparatus is found in Aristotle and dates back to about 1000 B.C.
. 1918. Sphyraena barracuda; Its morphology, habits, and history. Papers Tortugas
Laboratory 12: 53-108.
Carnegie Institution of Washington Publication Number 252.
This article provides a general description of the great barracuda, Sphyraena barracuda
made at the Tortugas Marine Laboratory, based on local collections and an examination of
12 large individuals using length/weight measurements, color and markings, jaws and teeth,
internal organs, foods and feeding, and manners of breathing. Additional information is
presented on their habits, how they may be caught, and parasites. An interesting historical
side of the paper compiled from around the world includes verbatim quotes and
descriptions of their great size, ferocity, fossil forms, nomenclature, habitats, and food
poisoning in man. Accounts of their poisonous flesh in the West Indies date as far back as
1667. Largest sizes of West Indies individuals approach 8-10 feet in length, with some
highly "dubious" reports of specimens reaching sizes of 18-20 feet in length.
. 1913. Uterine gestation in the nurse shark, Ginglymostoma cirratum. Journal of the Elisha
Mitchell Scientific Society 29: 8.
Also, in Science, 1913, v.37, p.993.
The breeding habits and embryology of this shark were studied at the Tortugas Laboratory
in the summer of 1912. A brief account was published in the Year Book for 1912, p. 148-
150.
34
134. Halley, R. B. and R. P. Steinen. 1979. Groundwater observations on small carbonate islands of
southern Florida. In Guide to sedimentation for the Dry Tortugas. Compiler R. B. Halley,
p. 82-89. Tallahassee, Florida: South East Geological Society Publication.
Observations are reported on the unusual hydrology of Loggerhead Key, a sandy key in the
Dry Tortugas in comparison with observations on Cluett Key, a mud key which lies 200 km
NE of Loggerhead in western Florida Bay. The ground water of Loggerhead and Cluett
Keys differs significantly from the surrounding sea water, despite the relatively small size
of the island. Climate alone does not determine the character of these ground waters; for
example, Loggerhead Key is underlain by less saline ground water than Cluett Key despite
the fact that it receives less rainfall. Ground water under such small islands such as these is
formed from topography, sediment character, vegetation, and many more parameters that
are themselves interrelated. They conspire to form ground water that not only differs from
sea water, but also can react with the island sediments to change the character of the
ground water. In this manner, island ground waters serve as geologic agents, hastening the
alteration of marine carbonate sediments to limestone and dolomite.
135. Hanlon, R. T. and R. F. Hixon. 1986. Behavioral associations of coral reef fishes with the sea-
anemone Condylactis gigantea in the Dry Tortugas, Florida USA. Bulletin of Marine
Science 39, no. 1: 130-134.
Over 30 small West Indian reef fishes dwell within the tentacular sphere of anemones,
mainly to avoid predation. Most species swim carefully to avoid the stinging tentacles, but
some species also have a physiological adaptation (skin mucus alteration) that allows them
to be in full and vigorous contact with the tentacles in a manner similar to Indo-Pacific
anemonefishes such as Amphiprion, Dascyllus and Premnas. The authors report herein six
species of reef fishes that are facultative associates of the sea anemone Condylactis
gigantea (Weinland) in the Dry Tortugas Islands. The fishes were not found associated
with other anemones. One species, Labrisomus gobio, is a new record of a fish with both
the behavioral and physiological adaptations to dwell unharmed among the stinging
tentacles of Condylactis gigantea.
136. Hargitt, C. W. 1911. Cradactis variabilis: An apparently new Tortugan Actinian. Papers Tortugas
Laboratory 3: 49-53.
Carnegie Institution of Washington Publication Number 132.
The author believes this species of actinian has never before been described, and names it
variabilis. The specimens seem to have the capacity to move about more or less freely,
and the frond-like organs situated about the margin of the oral disk and outside the outer
cycloe of tentacles aid in such movement. The color is pale olivaceous-green to brownish;
tentacles somewhat lighter; foliose organs darker, even brownish, with flake-white pads.
The body is highly contractile, with a weak or diffused sphincter. The reproductive season
seems to be in the spring and early summer. The habitat is chiefly in holes, crevices, or
similar secluded places in the coral reefs or about the shoals where protection is available.
137. Harrington, B. A. and J. J. Dinsmore. 1975. Mortality of transient cattle egrets at Dry Tortugas,
Florida. Bird-Banding 46, no. 1: 7-14.
This article examines the idea presented by Browder (1973) that cattle egrets pass through
the Dry Tortugas with seasonal regularity, and that large numbers die after landing. This
study concludes that regular spring movement occurs with many egrets stopping at the
island, and that many of the egrets that stopped apparently died from starvation, especially
in late June and in early July. The mortality in 1968 was higher that in 1970.
138. Harris, J. E. 1937. The mechanical significance of the position and movements of the paired fins in
the Teleosti. Papers Tortugas Laboratory 31: 171-89 (issued Oct. 1936).
Carnegie Institution of Washington Publication Number 475.
35
In the course of the evolution of the modern teleostean fish, a series of fairly well-defined
changes has taken place in the body form and in the shape and position of the fins. The
present paper discusses the mechanical factors concerned in the evolution of the teleost
type of fish. A comparison of this type with the dogfish suggests that the development of
an air bladder has been the primary factor involved in the change in general body form.
The reduction in specific gravity of the fish, consequent upon this primary change, has
removed the need for a lift force on the body during free swimming. The asymmetrical
(heterocercal) tail has therefore disappeared. For the same reason, the pectoral fins are no
longer needed as elevating planes, and become free to move up toward the mid line of the
body to act as brakes in stopping and turning movements. The forward motion of the pelvic
fins is a mechanism for producing a balanced vertical force and a balanced pitching
moment. These fins are normally used in conjunction with the pectorals. The independent
movements of the pectoral fins are then discussed. All types of movement so far observed
are variations on a fundamental form, in which the metrachronal oscillation of the fin rays
generates an undulating fin surface. The observed variations in form of the fin beat can be
produced by varying the phase difference between the beat of successive rays, and also by
making the oscillation of the fin ray asymmetrical. The characteristics of the pectoral
musculature associated with such variations are pointed out, and illustrated by reference to
a number of fish types.
139. Hartman, C. G. 1931. The hypophysis of fishes. Carnegie Institution of Washington, Year Book 30:
381-82.
Studies on the influence of the hypophysis on menstruation and various forms of uterine
bleeding in sharks were carried out.
140. Hartmeyer, R. 1911. Polycistor (Eudistoma) mayeri nov. sp. from the Tortugas. Papers Tortugas
141.
Laboratory 3: 89-93.
Carnegie Institution of Washington Publication Number 132.
A new species Polycitor (Eudistoma) mayeri , a new ascidian collected in 1907 at the
Tortugas is described as the largest and most beautiful ascidian of the Tortugas. It was
collected in the deeper water of the Southwest Channel near Loggerhead Key, on sandy
bottoms, where it is abundant. The color is pale yellow, with a reddish or violet tint. From
the western Atlantic only five species of this genus have been described, and all of these
are mentioned by Van Name from the Bermudas, but all these species have four rows of
stigmata in the branchial sac and are in many other respects quite different from this
species.
. 1908. Reisebilder aus Westinidien mit besonderer Berucksichtigung der korallenbildungen.
Deutsch. Gessel. Fur Volkstumlich Natuirkunde or Same Title in Meereskunde Jahrg. 3,
Heft 2, 40 Pp 3, no. 2: 1-40. (No abstract available)
142. Harvey, E. N.. 1911. Effect of different temperatures on the medusae, Cassiopea, with special
reference to the rate of conduction of the nerve impulse. Papers Tortugas Laboratory 3:
27-39.
Carnegie Institution of Washington Publication Number 132.
During the summer of 1909 a study was made of the effects of water temperatures on the
nerves and muscle tissue of Cassiopea. Temperatures in the moat at Fort Jefferson ranged
from 27°C to approximately 32-33°C. Activity limits and thermal death points of nerve and
muscle were measured. It was found that nerve conduction rates fall off in rate with rise of
temperature to a definite maximum, similar to that for enzyme action and for other life
processes.
36
143.
144.
145.
. 1914. The relation between the rate of penetration of marine tissues by alkali and the
change in functional activity induced by the alkali. Papers Tortugas Laboratory 6: 131-46.
Carnegie Institution of Washington Publication Number 183.
The present study, made at Tortugas in the summer of 1911, is a continuation of
permeability investigations undertaken at Columbia University in 1910 to 1911. The
author's aim has been twofold. First, to compare the permeability of the cells and tissues of
salt-water organisms with those of fresh-water forms. Second, to determine the relation
between the rate of penetration of the alkali and the appearance of structural or functional
changes in the cell. The author thinks that the presence of a sufficient number of OH ions
within the egg may aid in breaking down the granules and that this breaking down increases
also the degree of swelling of the egg. Cytolysis in Holothuris appears to be largely of this
type, since NaOH enters before the increase in volume begins. From this point of view
both theories of cytolysis contain an element of truth. Swelling of marine eggs is due both
to an increase in permeability of the surface and also to the breakdown of lipoid or protein
granules within. The latter tends to increase the swelling pressure or the osmotic pressure
of the egg, but is secondary to the increase in permeability of the surface.
. 1921. Studies on bioluminescence XIII: Luminescence in the Coelenterates. Biological
Bulletin 61: 280-287.
(No abstract available).
. 1923. Studies on bioluminescence. XV. Electroreproduction of oxyluciferin. Journal of
General Physiology 5: 275-84.
This work was on the light-producing reaction in the luminous crustacean, Cypridina.
Oxyluciferin may be reduced to luciferin at cathodes when an electric current is passed
through the solution, or at cathodes formed by metal couples in solution, or at cathodes of
oxidation-reducation cells of the NaCl - Pt - Na)S type. It is also reduced at those metal
surfaces (Al, Mn, Zn and Cd) which liberate nascent hydrogen from water, although no
visible hydrogen gas separates from the surface. Molecular hydrogen does not reduce
oxyluciferin even though very finely divided, but will reduce oxyluciferin in contact with
palladium. Palladium has no reducing action except in the presence of hydrogen, and
apparently acts as a catalyst by virtue of some power of converting molecular into atomic
hydrogen. Conditions are described under which a continuous luminescence of luciferin
can be obtained. This luminescence may be used as a test for atomic hydrogen. It is
suggested that the steady luminescence of bacteria is due to continuous oxidation of
luciferin to oxyluciferin and reduction of oxyluciferin to luciferin in different parts of the
bacterial cell.
146. Hatai, S. 1916. Changes in the chemical composition of starving Cassiopea xamachana. Carnegie
147.
Institution of Washington, Year Book 15: 206-7.
Studies were conducted on chemical changes occurring in Cassiopea during starvation.
The constancy of water content suggests that Cassiopea is largely a jelly-like mass, and
remains so throughout its life. In contrast, mammalian body-water content varies by age.
. 1917. On the composition of Cassiopea xamachana and the changes in it after
starvation. Papers Tortugas Laboratory 11: 95-109.
Carnegie Institution of Washington Publication Number 251.
For this study eight freshly caught normal Cassiopea, having different body weights, were
subjected to starvation by placing the animal in filtered sea water. The results were as
follows: 1. In general the smaller Cassiopea loses relatively more weight than the larger. 2.
The percentage of water found in the entire body is nearly the same in all sizes of
Cassiopea. However, the values of water content in the starved appear to be slightly higher
than those found in the normal Cassiopea. 3. The nitrogen content of the entire body is
148.
3H)
higher in the smaller than in the larger Cassiopea. 4. The absolute amount of nitrogen
found in the starved Cassiopea is considerably higher than in the normal having the same
body weight. It was noted that although high when compared with the normal, equal in
weight to the starved animal, it is very low for the initial body weight of the starved animal.
This shows that the nitrogen also has been consumed during the period of starvation. 5.
The nitrogen contents for the different parts of the body are simular in their relations to
those found in the normal Cassiopea. 6. The loss in weight of the different parts is of such
a character that their proportions in the starved remain similar to those in the normal
Cassiopea.
. 1917. On the composition of the medusa Cassiopea xamachama. Proceedings of the
National Academy of Science 3: 22-24.
In this study, an examination was made of three different parts of Cassiopea, mouth-
organs, umbrella, and velar margin to determine whether starving specimens lose weight
uniformly, or whether the loss is dissimilar in the three parts. Results indicated that the
smaller Cassiopea loses relatively more weight than does the larger Cassiopea. The
percentage of water is similar through the entire body, the nitrogen content is higher in the
smaller than the larger individuals, and nitrogen is much higher in the starved Cassiopea
than in the normal specimen with the same body weight. Results are compared with
Mayer's experiments, which showed nitrogen loss to be constant during the entire period of
starvation. Differences may be due to the size of animals used in his studies, as larger
individuals show little variation in nitrogen loss, whereas small Cassiopeas show large
variations in nitrogen loss due to body size.
149. Hayes, F. R. 1932. Nitrogen in echinoid ontogeny. Carnegie Institution of Washington, Year Book
150.
31: 284-85.
The chemical embryology of the echinoid egg was investigated, as well as variations in two
sources of energy available in the egg: protein and lipins.
. 1934. Variation in size and in nitrogen requirements during early development of the sea-
urchin, Echinomtera lacunter. Papers Tortugas Laboratory 28: 181-93 (issued Mar. 1933).
Carnegie Institution of Washington Publication Number 435.
After the penetration of a spermatozoon, the developing echinoderm egg receives nothing
from the outside except water and salts, until the comparatively advanced larva begins to
eat. The morphological phenomena of ontogeny can be brought about only by the
expenditure of energy, which must come from materials already present in the egg at the
time of fertilization. The problems of chemical embryology include (a) a determination of
the amount of energy required to produce structural changes, and (b) an investigation of the
chemical transformations taking place. The work here reported deals with a certain phase
of the chemical embryology of a common tropical sea-urchin, Echinometra lacunter.
Studies of the first 24 hours of development of the eggs of this form were carried on during
the summer of 1932 at the Tortugas Marine Station of the Carnegie Institution. Eggs of the
sea-urchin, Echinometra lacunter, were concentrated with a hand centrifuge and then
diluted with 500 times their volume. Analyses of primary amino nitrogen groups and of
total nitrogen were made, and the ratio of the former to the latter calculated. From 4 hours
onward the ratio of primary amino groups to total nitrogen increases. This does not mean,
however, a synthesis of the former at the expense of the latter, but rather that in the
combustion which provides the developing embryo with energy, some source of
nitrogenous fuel other than NH) groups is being used. There is a marked loss in the
quantity of nitrogen per egg during the period of development succeeding the first four
hours. One million eggs contain some 13 milligrams of nitrogen, of which about 28 per
cent is in the form of NH) groups.
38
151. Heard, R. W. and D. G. Perlmutter. 1977. Description of Colomastix janiceae, new species. A
commensal amphipod (Gammaridea Colomastigidae) from the Florida Keys, U.S.A.
Proceedings of the Biological Society of Washington 90, no. 1: 30-42.
During November of 1968 and 1973 and June of 1970 more than 100 specimens of an
undescribed commensal amphipod belonging to the genus Colomastix Grube, 1861 were
collected from loggerhead sponges, Spheciospongia vesparia (Lamarck), in the lower
Florida Keys. Additional specimens of this new species, collected from Dry Tortugas,
Florida were borrowed from the Division of Crustacea of the U.S. National Museum of
Natural History for examination.
152. Helwig, E. R. 1933. Regeneration in Jotrochota birotulata (Porifera). Carnegie Institution of
Washington, Year Book 32: 271-73.
The development and formation of cells over time was examined, from cross-sections made
from the branches of the sponge, Jotrochota birotulata.
153. Hendee, E. C. 1931. Formed components and fertilization in egg of the sea-urchin Lytechinus
variegatus. Papers Tortugas Laboratory 27: 99-105.
Carnegie Institution of Washington Publication Number 413.
This investigation of the eggs of Lytechinus variegatus collected during the summer of
1925 at the Tortugas was undertaken to determine if any substance of the egg was involved
in fertilization. Certain cytoplamic substances (macrosomes, hyaloplasm, chondriosomes,
fat droplets, and extra-nuclear basophilic granules) were demonstrated both before and
after fertilization. Lipid granules, present in the mature unfertilized egg, disappeared upon
fertilization.
154. Hendrix, S. A. and R S. Braman. 1995. NOx variation in the southeastern Gulf of Mexico. Florida
Scientist 58, no. 3: 292-97.
An automated system capable of providing speciation and concentration information for
several atmospheric NOx compounds was used to obtain diurnal and location variation data
during a five-day research cruise in the southeastern Gulf of Mexico approximately one
mile west of Fort Jefferson, Dry Tortugas between May 18 and May 22, 1987. Speciation
of these nitrogen compounds was achieved by selective preconcentration onto a series of
chemically coated glass hollow tubes. Analysis was performed by thermally desorbing the
collected analytes into a chemiluminescence detector providing sub parts-per-billion level
determination.
155. Hess, W. N. 1937. Reactions to light in Ptychodera bahamensis. Papers Tortugas Laboratory 31:
77-86 (issued Aug. 1936).
Carnegie Institution of Washington Publication Number 475.
Little attention has been given to the study of light reactions in any of the Enteropneusta,
and nothing is known, apparently, concerning the distribution or even the existence of
photoreceptors in this important group of animals. The purpose of this investigation was to
continue work on reactions to light and the photoreceptors in animals, using at this time a
more highly evolved species than the earth-worm on which the earlier work was done.
During the study, Ptychodera bahemensis responded negatively to ordinary intensities of
light. The movements of Ptychodera, when exposed to light were slow and deliberate and
there was little evidence of trial and error movements. The entire surface of the body was
sensitive to light, the most sensitive regions being on the proboscis and collar. Removal of
different parts of the body involving the central nervous system caused little if any decrease
in the percentage of negative responses to light. The reaction time of the proboscis was
greatly increased when it was removed from the rest of the animal. This is taken to
indicate that the central nervous system functions to speed up responses greatly, but is not
essential for responses. Removal of the proboscis together with the basal peduncle makes
156.
39
it impossible for the animal to orient when stimulated by light. This would seem to suggest
that the peduncle contains a coordinating center for certain bodily movements, or that the
animal has been rendered incapable of orienting, due to removal of that portion of the body
containing most of the notochord.
. 1940. Regional photosensitivity and photoreceptors of Crangon armillatus and the spiny
lobster, Panulirus argus. Papers Tortugas Laboratory 32: 153-61 (issued Sept. 1940).
Carnegie Institution of Washington Publication Number 517.
Crayfish from which the eyes have been removed are sensitive to light in the region of the
sixth abdominal segment, but no responses occurred when other regions were illuminated.
The discovery that freshly molted Crangon armillatus are sensitive to light in other regions
of their bodies, in addition to the sixth abdominal segment, led to this investigation. Results
of this study indicated that Crangon armillatus is usually sensitive to light in many regions
of its body, irrespective of how much time has elapsed since the last molting period.
Freshly molted spiny lobsters (Panulirus argus) are sensitive to light in many regions of
their bodies. Old spiny lobsters, with hard exoskeletons, from which the eyes have been
removed are usually not sensitive to light of the intensity used in these experiments. The
margins of the uropods of freshly molted Crangon armillatus and spiny lobsters are not
sensitive to light, but the basal two-thirds of these appendages are sensitive to light. Adult
Crangon armillatus and recently molted spiny lobsters react when illuminated from above
after the sixth abdominal ganglion has been shielded by black cardboard and also after the
ventral nerve cord has been cut between the fifth and sixth abdominal segments. This
shows that photosensitivity in these eyeless animals is not limited to the sixth abdominal
ganglion. Newly hatched Crangon armillatus with normal eyes swim toward the light with
their caudal ends foremost irrespective of the number of abdominal segments that have
been removed. Crangon armillatus and spiny lobsters from which the eyes have been
removed do not usually orient to light, but respond by random movements. When their
bodies are heavily pigmented, or if they are in poor physical condition, they do not respond
at all. However, if they do respond their responses are usually much slower than those of
freshly molted animals in good physical condition. The sixth abdominal segment of these
eyeless spiny lobsters and crayfishes is the most photosensitive region of their bodies.
However, in Crangon armillatus and the American lobster Homarus americanus all the
abdominal segments appear to be equally sensitive to light. On the basis of regional
photosensitivity of the uropods, it seems probable that the cell bodies of the neurons which
connect with the peripheral spines are sensitive to light and hence function as
photoreceptors.
157. Hoffman, W., and Jr. and P. C. Patty W. B. Robertson. 1979. Short-eared owl on Bush Key, Dry
Tortugas, Florida. Florida Field Naturalist 7, no. 2: 29-30.
The short-eared owl (Asio flammeus) is an uncommon but regular winter visitor to Florida.
This record represents the second summer record of Asio flammeus in Florida, and the first
record for the Dry Tortugas. The authors suggested that the bird in question had been on
Bush Key for some time, subsisting on the abundant tern chicks.
158. Holmes, C. W. 1984. Carbonate fans in the Florida Straits. Society of Economic Paleontologists and
Mineralogists Annual Meeting (Abstracts) 1: 39.
No abstract available.
159. Hooker, D. 1911. Certain reactions to color in the young loggerhead turtle. Papers Tortugas
Laboratory 3: 69-76 ahd illustrations.
Carnegie Institution of Washington Publication Number 132.
During the summer of 1907 observations and a series of experiments were made on the
habits and early life history of young loggerhead turtles, which identified reactions to color
40
and geotropism as the determining factors for the causes of young hatchlings to reach the
water. Based on day/night experiments on Loggerhead Key, hatchlings did not orient
towards the sun or the odor of the water, but exhibited positive phototropism by responding
to large surfaces of light of low intensity. After entering the water, the animal swam out to
sea apparently attracted by the darker blue of the deeper water. Young turtles displayed
positive geotropism when all possible negative geotropic reactions had been exhausted.
160. Hopkins, D. L. Locomotion/physiology of marine amoebae. Carnegie Institution of Washington,
161.
162.
163.
Year Book.: ;
1929,v.28,286-288: 1930, v.29,335-337.
The chemical and physical factors in the locomotion of marine amoebae collected from the
tidal pools at Tortugas and cultured in the laboratory were examined. Relationships
between sea-water salt and locomotion were determined by concentration and dilution.
Highest rates of locomotion were found in normal sea-water and could be a useful criterion
in classification and determining physiological condition in amoebae.
Jaap, W. C. 1985. An epidemic zooxanthellae expulsion during 1983 in the Lower Florida Keys
coral reefs: hyperthermic etiology. Proceedings of the Fifth International Coral Reef
Symposium , 143-48. Moorea, French Polynesia: Antenne Museum-Ephé.
Extensive reef coral zooxanthellae expulsion occurred from Key Largo to Dry Tortugas,
Florida, during September 1983. Coral bleaching was intensive between Pelican Shoal and
Sand Key Reef off Key West. Coral discoloration extended to depths exceeding 14 m but
was especially severe in shallow (1-2 m) spur and groove habitats. Approximately 75-95%
of all Millepora complanata and Palythoa caribaeorum were bone white, but most
colonies remained viable. Affected M. complanata (bladed fire coral) retained the ability
to inflict pain from dactylzooid nematocysts. Some individuals (5 to 10%) had fine algal
growth indicating death on all or parts of their skeletons. Although 15 species of
cnidarians, principally Scleractinia, were affected, some species (Madracis mirabilis,
Porites porites, Montastraea cavernosa, Dendrogyra cylindrus ) appeared to be immune.
A quantitative sample at Eastern Sambo Reef on 6 October documented 11 species and 209
colonies; M. complanata comprised 32.5% of all colonies. Transmission
eletromicrographs did not reveal epidemic pathogenic organisms in affected coral tissues.
Warm, calm weather prior to the expulsion was conducive to elevated seawater
temperature. A seawater thermograph deployed off Marquesas Key recorded temperatures
of 32.3 degrees C. during the period.
. 1980. Stony coral community structure at Long Key Reef, Ft. Jefferson National
Monument, Dry Tortugas, Florida. (abs.). Florida Scientist 43 (Suppl. 1).
Stony coral populations at Long Key Reef were studied during summers of 1975-76 under
National Park Service sponsorship. Plotless line transects (13, 25 m L ) were sampled in
depths of 0.5-21.3 m. Abundance, cover, and diversity were greatest in depths greater than
8 m. Of 34 species encountered, only 23 were censused quantitatively. Montastraea
annularis contributed 20% of all colonies and 37% of cover. Species richness was highest
(11) on transects in 7.6-12.5 m depths. Shannon-Weiner diversity values H' log SUB-2
computed by transects for individual colonies ranged from 1.0-3.0. Pielou's eveness (J')
values ranged from 0.36-1.00. Community relationships based on Morisita index values
detected an assemblage dominated by M. annularis in 8-13 m and a M. cavernosa
community in 18-21 m depths. Temporal comparison using the Morisita index revealed
strong community stability during 1975-76.
Jaap, W. C. and J. Wheaton. 1992. Summary of preliminary results, long-term ecological coral reef
studies, Ft. Jefferson National Monument, Dry Tortugas. Prepared for the National Park
Service Workshop, 28-30 April 1992, Miami, Florida.
41
Coral reefs exist over time scales of thousands of years. Processes of change in the
geological-time context occur slowly, e.g. sea level change correlated with glacial and
interglacial periods. The etiology of change is often poorly understood. For example, in
1878 a perturbation identified as, "black water” decimated Acropora spp. at Dry
Tortugas. Determining what black water was may never be known. Long-term ecological
research seeks to uncover processes that occur slowly or in which effects lag years behind
the causes. In the absence of long-term research, serious misjudgments can occur in
attempts to manage the environment. The National Park Service was interested in
developing a reef resource monitoring plan for Dry Tortugas reefs and collaborated with
the Florida Marine Research Institution in a joint study of reef resources. The goals of
these studies included testing methods, acquiring a data base on coral reef benthic and fish
communities to better understand the etiology of change, and isolating natural from
anthropogenic changes. Five study sites were selected in 1989. Repetitive sampling was
executed as precisely as possible using several different methods. These methods included:
transect sampling, video sampling, quadrat sampling, photographic sampling, recruitment
sampling, and environmental sampling. Results indicated that eleven octocoral, 22
scleractinian, and one milleporan species were enumerated on transects, while quadrats
indicated 29 octocoral, 26 scleractinian and 1 milleporan species over the study's three year
duration. Octocorals were consistently most diverse at Pulaski Shoal (20-21 species). Only
42 of 212 plates recruited scleratinian corals. This yielded an average of 0.35 recruit per
plate. A total of 187 milleporan corals recruited to 212 plates for an average of 0.88
recruit per plate (34.9/m?). The only octocoral recruit recorded was the gorgonacean
Briareum asbestinum, whose common name is corky sea fingers.
164.
. 1994. Summary of preliminary results, long-term ecological coral reef studies, Ft. Jefferson
National Monument, Dry Tortugas. Bulletin of Marine Science 54, no. 3: 1-10.
Narrative same as in reference no. 163.
165. Jaap, W.C., J. L. Wheaton and K. B. Donnelly. 1990. Materials and methods to establish
multipurpose, sustained, ecological research stations on coral reefs at Dry Tortugas. Diving
for Science... 1990. Proceedings of the American Academy of Underwater Science, Tenth
Annual Science Diving Symposium, 193-203. American Academy of Underwater Sciences:
Costa Mesa, California.
Sustained research requires precise, repetitive data acquisition to accurately evaluate
patterns of change in species abundance and community structure. Permanent reference
markers are essential to resample stations over time. The methods described here use solid
markers from which several sampling devices can be deployed. A hydraulic drill is used to
core 18-in deep holes into rock. A square stainless steel stake is inset, aligned, and
cemented into each hole. Quadrats, photogrammetric and video apparatus, and recruitment
arrays are deployed on or in reference to the stakes. Transects are extended between
stakes. The method is suitable for coral reef and other hard-bottom investigations.
166. Jaap, W. C., J. L. Wheaton , K. B. Donnelly, B. J. Kojis and J. E. McKenna Jr. 1994. A three year
evaluation of community dynamics of corals at Ft. Jefferson National Monument, Dry
Tortugas, Florida. Bulletin of Marine Science 54, no. 3: 1077.
Narrative same as in reference no. 167.
167. Jaap, W.C., J. L. Wheaton, K. B. Donnelly, B. L. Kojis and J. E. McKenna Jr. 1993. A three-year
evaluation of community dynamics of corals at Ft. Jefferson National monument, Dry
Tortugas, Florida, USA. (abs.). Proceedings of the 7th International Coral Reef
Symposium, page 164. Guam: University of Guam.
A study to evaluate methods and begin a long-term ecological research program at Ft.
Jefferson was initiated at five reef sites in May 1989. Benthos was mapped and
42
photographed within quadrats (5 x 2.56 m per site). Attached biota and substrates were
measured along 20- to 25-m transects (3 per site). Recruitment arrays were constructed of
PVC pipe, flat stock, and ceramic tiles (10.8 x 10.8 cm) and were secured to the reference
stakes. A carriage-mounted video camera, suspended on cables between two "T" poles
secured to the stakes, was pushed the length of a transect. Results implied relative stability
of the reef communities over three years. Dominant biota as determined by abundance and
cover remained similar. Classification analyses of station time-series also corroborated
relative stability. Recruitment of Millepora, Octocorallia, and Scleratinia was variable;
most recruits were found in cryptic refuge. The heterogeneity, high-relief, and multi-
layered canopy of these coral reef habitats restricts the usefulness of medium and long
distance (>1.5m) photography and video. We conclude that multiple sampling methods are
superior to a single sampling procedure.
168. Jaap, W. C., W. G. Lyons, P. Dustan and J. C. Halas. 1989. Stony coral (Scleractinia and
Milleporina) community structure at Bird Key Reef, Ft. Jefferson National Monument, Dry
Tortugas, Florida. Florida Marine Research Publication 46: 1-31.
Stony coral community structure at Bird Key Reef was investigated during 1975 using 30
continuous 25-m line transects in depths of 0.5 to 21.3 m. Thirty-two species, 872 colonies,
and 198 cm of coral cover were sampled quantitatively. Most species, colonies, and live
coral cover occurred seaward of 8-m depths on spur and groove substrate. Montastrea
annularis, M. cavernosa, and Siderastrea siderea constituted more than 50% of all cover.
Species diversity (Shannon index, log sub (2)) ranged from 1.0 for individual transects.
Diversity and envenness values computed from cover data were generally lower than
values computed from abundance data, reflecting M. annularis dominance. Numerical
community classification (Czekanowski's quantitative coefficient) revealed three groups
and an ecotone, each related to depth and substrate: 9 transects in 1 to 6 m depths
dominated by Porites asteroides and Diploria clivosa; an ecotone of 6 transects in 5 to 6
m depths; 5 transects in 6 to 9 m depths dominated by S. siderea; and 10 transects in 8 to
21 m depths dominated by M. annularis.
169. Jacobs, M. H. 1914. Physiological studies on certain protozoan parasites of Diadema setosum.
Papers Tortugas Laboratory 6: 147-57.
Carnegie Institution of Washington Publication Number 183.
It has been shown by the author and others that different species of protozoa have certain
physiological characteristics, often almost as stiking as their morphological ones, and
which are probably of considerable significance in the interpretation of their habits of life
and their relation to their environment. It occurred to the author to test a series of forms
which naturally live under essentially the same environmental conditions, and which may
be assumed to have done so for many past generations, in order to see whether they show
greater likenesses than a number of forms selected at random, or whether each has
preserved its individuality in spite of the similarity of its environment. The general results
of the experiments performed show surprising differences in the resistance of the parasites
of Diadema to various unfavorable conditions. In some cases the most resistant form may
live several hundred times as long as the least resistant one. Comparing all of the results
obtained, it is therefore seen that the similar habit of life of the four forms in question has
not brought about physiological similarity except in certain adaptive characters which are a
sine qua non for continued existence in the same host (e.g. ability to resist the digestive
Juices of the latter, etc.). In other respects they are just as different as almost any four free-
living forms that might be selected and the evidence of these experiments shows that the
physiological characters of an organism are not merely the result of its environment, but
may be as fundamental and characteristic as its morphological ones.
43
170. Jefferson, J. P. J. Y. Porter and T. Moore. 1879. On the destruction of fish in the vicinity of the
Tortugas during the months of September and October 1878. Proceedings of the U.S.
National Museum, Smithsonian Institution Press 1: 244-46.
The information in this report is relative to the die-off of large numbers of fish due to a
black water event in the Gulf of Mexico during the months of September and October
1878.
171. Jennings, H. S. 1909. Behavior of sea-anemones. Journal of Experimental Zoology 2: 447-72.
The study of the behavior of sea anemones (Stoichactis helianthus and Aiptasia spp.) was
made at the Carnegie Research Laboratory, Dry Tortugas using specimens collected in the
shallow waters near Fort Jefferson. Changes in behavior due to varying states of
metabolism for S. helianthus were examined using red meat, crab hard parts and filter
paper as food. After satiation, food is rejected through various reactions dependent upon
internal processes. Descriptions of food ingestion are described. For Aiptasia spp.,
experiments suggested that when the animals were hungry, they took both red and filter
paper; when satiated they took neither. Other topics of study included climatization to
stimuli (light), and reactions modified as a result of past experiences of the organism.
Results are compared to other lower groups of animals.
172. Jindrich, V. 1972. "Biogenic buildups and carbonate sedimentation, Dry Tortugas reef complex,
Florida." Ph.D. Dissertation, Geology, State University of New York at Binghamton.
The Dry Tortugas, a horseshoe-shaped complex of carbonate banks and coral reefs, is
located at the southern terminus of the Florida limestone shelf. The complex rises to the
surface waters from a drowned Pleistocene surface that forms a circular platform having a
general depth of 17-21 m. Three basic biogenic buildups (facies) comprise the reef
complex: 1) detrital lagoonal bank, 2) Montastrea reef bank and, 3) Acropora palmata
reef. These facies lie adjacent to one another and are also present in vertical succession as
individual growth stages of varying thickness and lateral extent. A zone of Acropora
cervicornis is developed as a transition between the Montastrea and A. palmata growth
stages. The present organic assemblages and topography bear evidence of dominantly
lateral progradation and cumulative storm effects that are linked to the slow eustatic sea-
level rise for the past several millennia. Long-continued storm degradation is manifested
by 1) continuous removal of A. palmata and its replacement by storm-resistant coralline
algae and Millepora sp. to produce truncated rocky surfaces, 2) abundant reef rubble, 3)
erosion of spur-grooves, and 4) development of intertidal rubble reef flats. Sediments
ranging from cobble-sized rubble to medium silt are composed of Halimeda, coral and
mollusc grains; coralline algae and foraminifers are present in minor amounts. Variations
in texture and constituent particle composition are interpreted to be mainly a result of mode
of sediment transport and effect of grain shape. Broadly-defined grain size populations
produced by three modes of transport have characteristic assemblages of constituent
particles. The populations include a gravel-sized surface creep population, sand-sized
saltation population, and very fine sand-to silt-sized suspension population. Strong mixing
occurs between the gravel and sand population on the storm-degraded shoals, and between
the sand and silt population on the lagoon bottom. Sand flanking the reefs and reef banks
shows minimum mixing hence good degree of sorting. Incongruous mixtures of the in-
place fraction and varying proportions of the transported populations constitute detrital
lagoonal banks as a substrate stabilized by seagrass and coral growth. The gravel-sand and
sand-silt mixtures are related to deposition under highly variable energy conditions.
Variability in energy conditions does not cause strong population intermixing on beaches.
For the same reason, beach sediments show a high degree of sorting in all size grades from
cobbles to fine sand.
44
173. Jones, N. 1938. Investigations on ascidians. Carnegie Institution of Washington, Year Book 37: 84.
The summer of 1938 was devoted to the study of the structure, development, budding, and
colony formation of Ecteinascidia tortugensis, a new ascidian species. The small ascidian,
one of the commonest during the season, occurred in large numbers on the under sides of
rocks just below low-water mark on both Bush Key and Long Key. Data report provided
by Plough and Jones, pp 97-98.
174. Jones, R. D. 1991. An improved fluorescence method for the determination of nanomolar
concentrations of ammonium in natural waters. Limnology and Oceanography 36, no. 1:
814-19.
An improved fluorescence method is described for measuring nanomolar concentrations of
NH, in natural waters. This method is based on the conversion of NH, to NH; and
subsequent diffusion of NH; across a microporous hydrophobic Teflon membrane into a
flowing stream of 0-phthaldialdehyde reagent to produce a fluorescent adduct. The
product is detected fluorometrically with a lower detection limit of better than 1.5 nM. Up
to 30 determinations h-1 can be made. The method works well in freshwater or salt water.
Field tests of the method in the Dry Tortugas and Gulf Stream gave NH, concentrations
that ranged from 18.0 nM in Gulf Stream waters to 2,254.7 nM in interstitial waters from
coralline reef sands. The method can be used to measure near real-time NH,
concentrations in situations where it was previously difficult or impossible.
175. Jones, R. S. and M. J. Thompson. 1978. Comparison of Florida reef fish assemblages using a rapid
visual technique. Bulletin of Marine Science 28, no. 1: 159-72.
Species composition, species diversity, and relative abundance of 4 coral reef fish
communities in John Pennekamp State Park, Key Largo, Florida, are compared with 4
communities at Fort Jefferson National Monument in the Dry Tortugas using the species-
time, random-count technique. The technique is similar to species-area methods, but time
replaces area. Fish communities at Pennekamp Park showed the highest overall number of
species and scores (reflecting species abundance, and species diversity). Two artificial
reefs (shipwrecks) included in the study both show closer relationships to adjacent reefs
than to wreck-specific species.
176. Jordan, D.S. 1904. Notes on fishes collected in the Tortugas Archipelago (by Dr. Joseph C.
Thompson). Bulletin of the United States Fish Commission for 1902 22: 539-44.
An additional sixteen species of fish are described for the Tortugas based on collections
made by J.C. Thompson while on the northward cruise of the steamer Chesapeake.
177. Jordan, D. S. and J. C. Thompson. 1905. The fish fauna of the Tortugas Archipelago. Bulletin of the
United States Bureau of Fisheries for 1904 24: 229-56.
The shallow water fishes of the Tortugas, as suggested by A.G. Mayer to David Starr
Jordan, are unsurpassed in variety and abundance anywhere along the Atlantic coast of the
United States, and based on the nearness of the Gulf Stream and the winds and currents,
pelagic fish from all over the Gulf of Mexico and the West Indies may be drifted by the
Tortugas. Collections made by Thompson while on duty as a medical doctor at the Garden
Key Naval Station resulted in an annotated fish list containing 218 species known to occur
at the Dry Tortugas at the time.
178. Jordan, H. E. 1908. The accessory chromosome in Aplopus mayeri. Anatomischer Anzeiger Bd 32:
284-95.
The purpose of this paper is to trace the accessory chromosome in the phasmid, Aplopus
mayeri from material collected from Loggerhead key, Florida. The accessory chromosome
appears in the resting stage of the secondary spermatogonia as a chromatin nucleolus
characteristically close to the nuclear wall. Both the primary and secondary spermatogonia
72),
180.
181.
182.
45
have a metaphase group of 35 chromosomes. The accessory chromosome can be traced as
a specific structure from the resting stage of the last order of spermatogonia through all the
various phases of synapsis and maturation, until it disintegrates in the head of the ripening
spermatozoa.
. 1917. Aortic cell clusters in vertebrate embryos. Proceedings of the National Academy of
Science 3: 149-56.
Aortic cell clusters are described among various animal groups (pig and chick) and
compared to 12-day loggerhead turtle embryos and mongoose embryos. Various aspects of
the hemogenic activity of embryonic endothelium are described consequent to the inherent
capacity of endothelium to produce hemoblast, and not in connection with an associated
toxic substance.
. 1917. Atresia of the esophagus in the embryo of the loggerhead turtle, Caretta caretta: A
normal developmental condition. Papers Tortugas Laboratory 11: 345-60.
Carnegie Institution of Washington Publication Number 251.
A series of 26 embryos of the loggerhead turtle were collected and used originally for a
study of the history of the primordial germ-cells. It was noticed that the esophagus was
solid for a greater or less extent, approximately from the point of origin of the respiratory
anlage to its bifurcation into the bronchi, from the eleventh to the thirty-second day of
incubation. Points of special significance in regard to this material are: (1) the relatively
longer persistence of the occlusion than has yet been described for any other form; (2) the
absence of contributory yolk in the stenosed area; (3) close relation to the point of origin of
the respiratory anlage, which fact may disclose its possible functional significance .
. 1917. Embryonic history of the germ-cells of the loggerhead turtle (Caretta caretta).
Papers Tortugas Laboratory 11: 313-44.
Carnegie Institution of Washington Publication Number 251.
The wide discrepancies in the published accounts of the origin and early history of the
germ-cells in vertebrates provided the stimulus for this investigation. Twenty-five embryos
of the loggerhead turtle (Caretta caretta), ranging from the second day (5 somites, 2mm.
length) to the thirty-second day of incubation, were employed in this investigation. Results
indicate that the primordial germ-cells migrate during the second day from the yolk-sac
endoderm, where they are widely scattered caudally, into the lateral border of the area
pellucida on each side of the embryonic disk. The germ-cells migrate by amoeboid
activity. The migration period is not sharply limited. A certain number of germ-cells
migrate out of the regular germ-cell route and go astray. The total number of primordial
germ-cells counted in a 12-day embryo is 352. Occasional cells may divide by mitosis, or
undergo degeneration, at any stage of their history or at any point of the route. No germ-
cells were found contributing to the formation of the Wolffian duct. The germ-cells do not
differ from young somatic cells in the character of their mitochondrial content. No
transition stages between coelomic epithelial cells and germ-cells appear up to the 32-day
stage. The evidence derived from a study of the Caretta embryos is in complete harmony
with the idea of a single uninterrupted line of sex-cells from primordial germ-cells to
odgonia and spermogonia, and with the hypothesis of a vertebrate Keimbahn or continuous
germinal path.
. 1908. The germinal spot in echinoderm eggs. Papers Tortugas Laboratory 1: 1-12.
Carnegie Institution of Washington Publication Number 102.
This paper reports the results of further studies of the prematuration stages of echinoderm
eggs of additional species of echinoderms, a star-fish (Echinaster crassispina), and a
brittle-star (Ophiocoma pumila). In Echinaster crassispina the chromosomes are derived
exclusively from the nucleolus. In Ophiocoma pumila the chromosomes arise exclusively
46
183.
184.
185.
from the nuclear reticulum. In some species the chromosomes arise from a chromatin-
nucleolus, in others from a chromatic reticulum, and in still others in part from one source
and in part from the other. The eggs of different forms differ in that some have only a
chromatin-nuleolus, without distinct plastin ground-substance, resting in an achromatic
nuclear reticulum (Echinaster); others possess both chromatin-nucleolus and plasmosome
as well as a chromatic nuclear reticulum (Ophiocoma); and still others possess a double
nucleolus (chromatin nucleolus and plastin ground-substance), with the chromosome
complex gathered in a mass in the achromatic reticulum (Asterias). The function of the
germinal spot then appears, in part at least, to be that of a storehouse of material which is to
contribute to the formation of the chromosomes .
. 1917. The history of the primordial germ cells in the loggerhead turtle. Proceedings of the
National Academy of Science 3: 271-75.
This study attempts to trace germ cell history in the loggerhead turtle and compare it to
observations for other vertebrates. Embryos were collected for study from specimens taken
on Loggerhead Key, Dry Tortugas during the Summer of 1914. The germ cell history of
Caretta is very similar to that first described for Chrysemys and to that described for
dogfish.
. 1917. The microscopic structure of striped muscle of Limulus. Papers Tortugas Laboratory
11: 273-90.
Carnegie Institution of Washington Publication Number 251.
The study of the skeletal muscles of Limulus was undertaken with two chief objects in
view: to test a conclusion suggested by earlier studies on the intercalated disks of
vertebrate cardiac muscle, namely, that these disks are properly interpreted as "irreversible
contraction bands" and to seek additional evidence in further refutation of the recently
revived hypothesis that striped muscle can be interpreted in terms of "muscle-cells" and
intercellular myofibrillae. It was found that both the skeletal and the cardiac muscles of
Limulus consist of trabeculae of finely granular sarcoplam. In cardiac muscle the main
trabeculae and their branches form a loose-meshed syncytium. Neither type of muscle
contains mesophragmata. Very rarely an intercalated disk of the simple-comb type appears
in the cardiac muscle. Both types are very similar in respect of the presence and
arrangement, in the same phase of contraciton, of Q and J disks, and the telophragmata.
The evidence is unequivocal against an interpretation of structure in terms of "muscle-
cells" and intercellular myobibrillae. The nuclei of the growing muscles multiply by
amitotic division. In essential structure the cardiac and skeletal muscles of Limulus are
closely similar, indicating a close functional similarity. The structure serves, moreover, as
a splendid illustration of the "law of biogenesis," in that it is practically identical with a
stage in the early histogenesis of striped muscle of teleosts .
. 1908. The relation of the nucleolus to the chromosomes in the primary odcyte of Asterias
forbesii. Papers Tortugas Laboratory 1: 37-72.
Carnegie Institution of Washington Publication Number 102.
The primary object of this investigation was was to contribute to the subject of the relation
between nucleolus and chromosomes during maturation. In summary, synizesis occurs in
the odcyte of the first order at the very beginning of the growth-period (size of nucleus 5
microns). The growth-period is passed through rapidly. During the latter half of the
growth-period all the chromatin, with the exception of what is held by the chromosomes,
becomes stored in the enlarging nucleolus. The nucleolus consists of a plastin ground-
substance infiltrated and covered over with chromatin. The chromosomes do not arise out
of the nucleolus. The number of chromosomes in the prophase of the first polar mitosis is
18. They vary somewhat in size (one is considerably larger than the rest), all have a
characteristic dumb-bell shaped appearance, and some are clearly double (bivalent). The
47
two maturation divisions effect a double longitudinal fission of the original bilobed
chromosomes. The reduced number of chromosomes is again 18. Observations on
Hipponoé esculenta agree in essential points with those made on Asteria forbesii and
support the conclusions regarding the origin of the chromosomes, the function of the
nucleolus, and the reduction phenomena.
186.
. 1908. The spermatogenesis of Aplopus mayeri. Papers Tortugas Laboratory 1: 13-36.
Carnegie Institution of Washington Publication Number 102.
The object of the present investigation is primarily to trace the history of the accessory
chromosome through the various stages in the process of spermatogenesis in the phasmid
Aplopus mayeri. The material upon which the investigation is based was obtained from the
Loggerhead Key, Florida. Primary spermatogonia divide both mitotically and amitotically.
In the latter instance cell-division is frequently not consummated and a bi- or multi-nuclear
cell results. In the first order of the secondary spermatogonia the accessory chromosome
appears in the resting-stage. During synapsis the accessory chromosome lengthens into a
club-shaped structure attached by its lesser end to the presynaptic thread, undergoes partial
longitudinal division, closes up again during the height of synapsis, and returns again to its
previous characteristic form and location in the nucleus of the growing primary
spermatocyte. The second maturation division is equational, effecting a longitudinal
division of univalent chromosomes. The accessory also divides equationally in the cells
containing this element. A dimorphism of spermatozoa results; the accessory chromosome
possessed by one-half probably represents a sex-determinant. The history of the accessory
chromosome gives evidence that it at least possesses a strict morphological and probably
also a physiological individuality.
187. Kaas, P. 1972. Polyplacophora of the Caribbean region. P. Wagenaar and L. J. Van Der Steen
Hummelinck, 1-162. Studies on the Fauna of Curacao and Other Caribbean Islands, ed. P.
Wagenaar and L. J. Van Der Steen Hummelinck. The Hague: Martinus Nijhoff.
This paper includes Tables of Distribution of Polyplacophora of the Caribbean . The
author took into consideration the whole of the Florida coast as far north as Fernandina, E.
Florida and the Keys, the Dry Tortugas, W. Florida, the Gulf of Mexico, and also the
Bermudas. Thiele's description of his single 5.5 specimen of Ischnochiton hartmeyeri from
Bird Key Reef is translated into English here by Kaas.
188. Kale, H. W. 1985. Florida birds - Dry Tortugas. Florida Naturalist 58, no. 2: 6.
A sighting of a great black-backed gull is made at the Dry Tortugas, and a scarcity of land
birds is reported.
189. Kellner, Carl. 1907. Embryology of the appendicularian, Oikopleura. Zoological Anzeiger, Bd. 31:
May.
The appendicularia of the Dry Tortugas specimens of Salpae were collected.
Appendicularia of the genus Oikopleura and their "houses" were examined and found in
surface waters. Their anatomy and histology are described.
190. Kille, F. R. 1936-1937. Regeneration in holothurians. Carnegie Institution of Washington, Year
Book.
Note: published as follows: 1936, v. 35, p. 85-86; 1937, v. 36, p. 93-94.
Histological studies were conducted on sea-cucumbers of the genus Holothuria, to
determine the manner in which the digestive system is reconstituted following autotomy by
means of electrical stimuli.
191. Kopac, M. J. 1936. Electrical resistance of Valonia. I. Changes in the resistance with time in
impaled coenocytes. Papers Tortugas Laboratory 29: 359-86 (issued Mar. 1936).
48
Carnegie Institution of Washington Publication Number 452.
During the summers of 1933 and 1934 the author worked at the laboratory of the Carnegie
Institution located on Loggerhead Key, Dry Tortugas, Florida. Several species of Valonia
were found growing abundantly on the various coral reefs of the Dry Tortugas. A study of
the electrical resistance of impaled Valonia coenocytes by using a technique more highly
refined that that employed by previous investigators was initiated. Although only V.
ventricosa was used in this study, the methods developed and used here may be extended
to the study of other species of Valonia. Glass microcapillaries, with tips ranging from
0.025 to 0.1 mm. in diameter and filled with vacuolar sap, served as microsaltbridges
leading from the vacuole to a calomel half-cell. A larger glass tube (the macrosaltbridge),
filled with sea-water, was used as a saltbridge leading from the sea-water surrounding the
coenocyte to another calomel half-cell. The Valonia coenocytes were impaled on the tip of
the microsaltbridge with the aid of a micromanipulator. It was found that coenocytes with a
high chloroplastid density have a constant Rp several times higher than coenocytes with a
low chloroplastid density. It is postulated that only the inter-chloroplastidal protoplasm is
capable of conducting a current. The average initial Rp in type A punctures was 60 to 65
per cent of the constant Rp. In some coenocytes a constant Rp was reached in a few
minutes. This increase in Rp is caused largely be the redistribution of those chloroplastids
around the microtip which were disturbed by the puncture. The average initial Rp in type
B punctures was 2 to 3 per cent of the constant Rp. After the chloroplastids are
redistributed in this hyaline zone, the disintegrated chloroplastids are extruded, and the tiny
vauoles are eliminated, no further increase in Rp takes place .
192. Kunkel, B. W. 1934. The selective action of certain adverse environmental conditions on the hermit
crab (Clibanarius tricolor Gibbes). Papers Tortugas Laboratory 28: 215-44 (issued Aug.
1933).
Carnegie Institution of Washington Publication Number 435.
The problem of selection is undoubtedly a very complex one. The characters which enable
one organism rather than another to survive are difficult to ascertain; a favorable variation
of one part may be accompanied by an unfavorable variation of another, so that selection
may have no effect upon the first feature. The present study has to do with a phase of the
selection problem which, on the whole, has received rather scant attention from
investigators. The selective effect of certain adverse conditions on a population has been
studied. The problem is that of determining how a given species may respond to a change
in environment, of determining the morphological difference between those individuals
which succumb to a certain change in the normal environment and those which are able to
withstand the change. The material upon which the present study is based was collected
and the experiments were made at the Tortugas Laboratory. The small hermit crab
Clibanarius tricolor Gibbes was selected for the experiments.
193. LeCompte, M. 1937. Some observations on the coral reefs of the Tortugas. Carnegie Institution of
Washington, Year Book 36: 96-97.
Particular attention was paid to the distribution and adaptation of the corals on the reef
west of Loggerhead Key. A baseline of about 2500 yards is verified, documenting large
heads of Orbicella (Madrepora) annularis, extensive growths of Acropora, and areas of
gorgonians and algae. Beach rock development, coral feeding habits, and the effects of
boring animals on corals are discussed.
194. Leitch, James L. 1936. The water exchanges of living cells. II. The application of a photographic
method to the determination of the non-solvent volume of the eggs of Echinometra
lacunter. Papers Tortugas Laboratory 29: 349-58 (issued Mar. 1936).
Carnegie Institution of Washington Publication Number 452.
Photography has been applied to the study of living cells and tissues. Leitch raised the
49
question of the feasibility of a photographic method in the study of the osmotic behavior of
cells. The present paper outlines such a method and discusses some of the factors involved
in the study of the water exchanges of the eggs of the sea-urchin, Echinometra lacunter,
using measurements of photographs of eggs at equilibrium in dilute sea-water solutions. It
was shown that photography can be employed in the study of the water exchanges of living
cells. The non-solvent volume of the eggs of Echinometra lucunter is 36 per cent when
calculated after from 60 to 90 minutes’ exposure to experimental solutions. Longer
exposures to the experimental solutions result in a higher non-solvent volume of 48 per
cent which is associated with pronounced vacuole formation. The appearance of vacuoles
after the attainment of the first equilibrium is discussed and three different explanations
proposed.
. 1937. The water exchanges of living cells. IV. Further studies on the water relations of the
eggs of the sea-urchin, Echinometra lacunter. Papers Tortugas Laboratory 31: 53-70
(issued July, 1936).
Carnegie Institution of Washington Publication Number 475.
The application of a photographic method to the determination of the non-solvent volume
of the eggs of the sea-urchin, Echinometra lacunter, and also the effect on the non-solvent
volume determinations of the length of time of immersion of eggs in diluted sea-water
solutions, was demonstrated in another publication (Leitch,1936). The present paper
considers the utilization of this photographic method for the study of the swelling and
shrinking of the eggs of the same sea-urchin, the effect on the water relations of these cells
of the time between that of spawning and that of introducing the eggs into the experimental
solutions, and an analysis of equations which have been developed and applied by several
investigators to explain the kinetics of water exchanges of living cells. Results showed that
an analysis of the equations developed to interpret the kinetics of water exchanges of cells
the so-called permeability "constants" are not constant for the eggs of Echinometra
lacunter but vary with the dilutions of sea-water used and also with different intervals of
time in the same dilution. The permeability constants for swelling and shrinking do not
coincide, being between 0.250 and 0.650 for the former process and between 0.180 and
0.580 for the latter. The introduction of the correction for the non-solvent volume into the
equations does not produce a better agreement between the constants for the two processes.
The use of the photographic method (Leitch, 1936) is further substantiated for the
determination of non-solvent volumes and is extended to the study of the swelling and
shrinking of eggs. Approximately two hours from the time of spawning the non-solvent
volumes is greatly increased, from 30 to 53 per cent of the initial volume. There is a slight
retarding effect on the water exchanges of the eggs brought about by standing. The time at
which the effect of standing at room temperature appears in the values of the non-solvent
volume and rate of penetration of water is correlated with a sharp decrease in the
percentage of development, a slight increase in the volume of the eggs and a cytolysis-like
phenomenon which finally ends with the complete disintegration of the eggs. The
production of fertilization membranes as a criterion of non-injury of the egg cell is shown
to be inadequate and the percentage of development of normal larvae is urged as a better
test of normality .
195.
196. Lessios, H. A., D. R. Robertson and J. D. Cubit. 1984. Spread of Diadema mass mortality through
the Caribbean. Science 226: 335-37.
Populations of the ecologically important sea urchin Diadema antillarum suffered severe
mass mortalities throughout the Caribbean. This mortality was first observed at Panama in
January 1983; by January 1984 it had spread to the rest of the Caribbean and to Bermuda.
The sequence of mortality events in most areas is consistent with the hypothesis that the
causative agent was dispersed by major surface currents over large distances. However,
some of the late die-offs in the southeastern Caribbean do not fit this pattern. Several lines
50
of indirect evidence suggest that the phenomenon is due to a water-borne pathogen. If so,
this is the most extensive epidemic documented for a marine invertebrate.
197. Linton, E. 1908. Helminth fauna of the Dry Tortugas. I. Cestodes. Papers Tortugas Laboratory 1:
198.
199.
200. Lipman,
201. ———.
157-90.
Carnegie Institution of Washington Publication Number 102.
This report is based on data collected at the Marine Biological Laboratory, Tortugas,
Florida, June 30 to July 18, 1906. A list of the hosts which were examined for parasites,
and a summary of the results of that examination, together with a few food notes are
presented. A few extracts from notes made at the time the material was collected are
presented. Acanthocephala are presented. The species found in the frigate mackerel was
Echinorphynchus pristis. Few nematodes were found in the nurse-shark. New species of
parasites are described.
. 1910. Helminth fauna of the Dry Tortugas. II. Trematodes. Papers Tortugas Laboratory 4:
11-98.
Carnegie Institution of Washington Publication Number 133.
The collection here described was made at the Marine Biological Laboratory, Tortugas,
Florida, in the summers of 1906, 1907, and 1908. The fishes examined were from the
shallow waters of the reef. The distribution of parasites together with food notes have
already been published in the Year Book of the Carnegie Institution of Washington for the
years above named. This paper includes a list of Tortugas trematodes and their hosts, key
to the genera and species described, and descriptions of species, habitats, etc, including:
trematodes from loggerhead turtles and from fish.
. 1907. Note on the habits of Fierasfer affinis. American Naturalist 41, no. 481: 1-4.
Observations of the Fierasfer affinis entering its host, tail first are made.
C. B. 1929. The chemical composition of sea water. Papers Tortugas Laboratory 26: 249-
Vile
Carnegie Institution of Washington Publication Number 391.
In his studies on marine bacteria and related subjects, the author realized the need for more
accurate and complete analytical data on the inorganic components of sea-water and
determined to obtain them.. Two samples were analyzed from the Atlantic area and they
were both from the Gulf Stream, and taken near Loggerhead Key in the Tortugas. The data
render it clear that a large part of the ions important to algae are removed from solution in
sea-water by a rise in pH of that medium, which is well within the range of daily rise in pH
of sea-water carrying an active algal flora under the proper conditions of light and
temperature.
1924. A critical and experimental study of Drew's bacterial hypothesis on CaCO;
precipitation in the sea. Papers Tortugas Laboratory 19: 179-91.
Carnegie Institution of Washington Publication Number 340.
Based on a series of experiments to explain the precipitation of CaCO; in sea water, it was
found that there are several ways to explain CaCO3, where it occurs in seawater, without
introducing Drew's hypothesis or any other bacterial hypothesis. These explanations of the
phenomenon seem adequate to account for the qualitative and quantitative differences in
CaCO; as found under different conditions in seawater. Changes in water and air
temperatures, and marine plant activity, which Drew and others clearly appreciated and
understood, yet have introduced a purely gratuitous bacterial hypothesis based on what
appears to be sound experiments.
51
202. . 1929. Further studies on marine bacteria with special reference to the Drew hypothesis on
CaCo; precipitation in the sea. Papers Tortugas Laboratory 26: 231-48.
Carnegie Institution of Washington Publication Number 391.
Bacterial populations in the open sea are very small. Mixed or pure cultures of organisms
isolated from sea-water are incapable of precipitating CaCO; from sea-water to which no
salts have been added, or even in the presence of added KNO3 Mixed or pure cultures of
such organisms are incapable of precipitating CaCO; in a sea-water medium if KNO; and
organic matter as the sugars or similar forms free from calcium are added to the medium.
Upon the basis of evidence in this and in earlier papers the Drew hypothesis is shown to be
untenable, and at the very least uproved. This strong probability is reemphasized in the
purely physical-chemical nature of CaCO; precipitation on a large scale in nature.
203. Locker, S. D. A. C. Hine and E. A. Shinn. 1991. Sea level geostrophic current control on carbonate
shelf-slope depositional sequences and erosional patterns, South Florida platform. AAPG
Bulletin 75, no. 3: 623.
High-resolution seismic reflection profiles across the shelf-slope margin between the Dry
Tortugas and Key West, Florida, indicate that sea-level fluctuations and the eastward
flowing Florida Current are major controls on late Quaternary sequence stratigraphy. The
study area, a transition zone between the open south Florida shelf and the lower Florida
Keys island/reef system, is typified by a shallow shelf with reef margin adjacent to a deeper
lower-shelf/slope. The lower-shelf/slope is composed of stacked or prograding sequences
that downlap and pinchout on the Pourtales Terrace. Strike oriented stratigraphic sections
exhibit many sea-level controlled features such as lowstand erosion, transgressive
unconformities, and highstand system tracs. Lowstand reefs, notches, or barriers are
observed as deep as 150m below present sea level. Depositional styles change along-slope
from west to east. The western portion of the study area is characterized by thick, low
amplitude prograding sequences related to abundant supply of sediment through off-shelf
transport during high sea-levels as well as along-slope reworking by the Florida current.
Part of this section has been severely eroded by along-slope current producing localized
cur and fill structures and widespread erosional unconformities. To the east, a thinner
section of high-amplitude reflections is common seaward of the lower Florida Keys reef
tract system. Again, along-slope current erosion and winnowing of sediment supplied by
the adjacent margin is evident. This study provides new evidence of how a strong
geostrophic boundary current along with fluctuating sea levels have interacted to control
depositional sequences on a carbonate slope in the Florida/Bahamas platform complex.
204. Longley, W. H. 1917. Changeable coloration in Brachyura. Proceedings of the National Academy of
Science 3: 609-11.
Studies on changes in the color of brachyuran crabs (Ocypoda and Callinectes sp.) at the
Marine Laboratory, and in the field on Loggerhead Key, Dry Tortugas, demonstrate
adaptive coloration based on temperature variations and the color of the substratum upon
which the specimen is resting. It is expected that in future studies the same general rules of
adaptation for fishes will apply to crabs.
205. . 1918. Haunts and habits of tropical fishes. American Museum Journal 18: 79-88.
Observations are recorded at the Dry Tortugas using underwater photography. Habits of
the shallow water reef species were photographed in water less than 10 feet deep.
Emphasis is placed on the biological significance of color in fishes with their surroundings.
Fish color change may be evoked by offering them food by hand at different locations.
The foods and feeding habits of reef fish are discussed in this report.
206. ————. 1918. Marine camoufleurs and their camouflage: the present and prospective significance
of facts regarding coloration of tropical fishes. Smithsonian Report (1920): 475-85.
a2
207.
208.
209.
Fish are used as an example of an animal which uses color patterns, not as a struggle for
existence as hypothesized by Darwin, but as a means of expressing its biological
significance by displaying a natural system of camouflage. Some colors in fishes are not
changeable, but seem to be correlated to definite habits. In the case of those that are
changeable, there is conclusive evidence that they are displayed under specific conditions.
For example, transverse bands are shown when the species is inactive. However, upon
movement the bands are replaced by stripes.
. 1916. Observations upon tropical fishes and inferences from their adaptive coloration.
Proceedings of the National Academy of Science 2: 733-37.
The conception that species have been multiplied by divergent evolution of related strains
is based on many observations. If the Darwinian hypothesis is true, the character of
organisms should be largely of an adaptive sort, but its adherents have been content to
support this position by inputting utility to structure and habits. It has not been proved in
fishes that some color combinations ward off enemies nor that pigmentation is functionally
conspicuous. Many of the brightly colored fishes of the Tortugas have been studied to
evaluate their coloration objectively. Most species exhibit countershading with darkest
shading on the upper surface and lighter shading on the mid-ventral or lower line. Thirteen
species of fish studied exhibit color changes based on their surroundings observed from
boats or from the bottom using diving equipment and photography. Correlation of color
with habitat has been documented. Some examples suggest that red fish are rarely seen
during the day, gray fish with diurnal activity patterns are found near large coral heads,
lighter blue fish are habitually found swimming well above the bottom in moderate depths,
and those species largely found over grass beds are of green color or have a green color
phase. As far as this class of animals is concerned, Longley postulates that there is no
ground for the belief that bright color is correlated any way with armament or
distastefulness. Problems of mimicry resemblance are unresolved, however the
observations presented in this abstract undermine many speculative explanations of animal
coloration in terms of natural selection and replace them with something which may not be
dismissed from consideration.
. 1936. Species studies and the species problem. American Naturalist 70: 97-109.
(No abstract available).
. 1917. Studies upon the biological significance of animal coloration. I: The colors and color
changes of West Indian reef fishes. Journal of Experimental Zoology 23: 536-601.
Studies were carried out at the Dry Tortugas to determine the biological significance of
changes in color of reef fishes. It was found that fishes are countershaded; color changes,
which are common even among the most gaudy, tend to assimilate them with their
environment; and in general, their colors repeat those of their surroundings. Specially
defended types are not unlike others in pigmentation, nor inferior to them in their ability to
effect adaptive color adjustments. Finally, there is no evidence that brightly colored
species enjoy greater immunity from attack than their fellows, for they constitute a large
proportion of the food and may be readily identified in the stomach contents of predaceous
forms. These statements, which rest upon a great body of verifiable observations, are
consistent with the Darwinian hypothesis, but inconsistent with the assumption that animals
of high color possess more than minimal conspicuousness under natural conditions. They
impel one to reject the hypotheses of warning and immunity coloration, signal and
recognition marks, and sexual selection, at least in so far as they may ever have been
supposed to apply to these forms. Upon the contrary, they confirm Thayer's conclusions
regarding the obliterative function of color and pattern, emphasize the common occurrence
of adaptive characters among animals, and suggest that their evolution has been guided
throughout by natural selection.
53
210. . 1917. Studies upon the biological significance of animal coloration. II: A revisional
working hypothesis of mimicry. American Naturalist 51: 257-85.
In this report, various hypotheses proposed by the author and other investigators relating
changes in animal coloration in relation to habits are discussed. The author postulates that
bright colors of tropical fishes are correlated with the animal's habits from work achieved
at the Dry Tortugas. Other coloration hypotheses are provided dealing with butterflies and
lizards, as well as warning coloration in bright and dull-colored insects. These ideas
submitted by the author constitute working hypotheses to be tested by other biologists.
211. Longley, W. H. and S. F. Hildebrand. 1940. New genera and species of fishes from the Tortugas,
Florida. Papers Tortugas Laboratory 32: 223-85 (issued Sept. 1940).
Carnegie Institution of Washington Publication Number 517.
Thirty new genera and species of fishes described in these pages resulted from studies
carried on for many years, principally at Tortugas, Florida, by the late Dr. William H.
Longley, whose untimely death occurred before he had fully completed a study of his
collections and a manuscript embodying a complete account of his field observations. The
present writer has made further studies of the Tortugas collections, and has endeavored to
extract interesting facts from Dr. Longley's notes on those species not treated in his
unfinished manuscript.
212. Longley, W. H. and S. F. Hildebrand. 1941. Systematic catalogue of the fishes of Tortugas, Florida;
with observations on color, habits, and local distribution. Papers Tortugas Laboratory 34:
1-331.
Carnegie Institution of Washington Publication Number 535.
Observations on the fishes of the Tortugas Atoll were made by the senior author spanning a
period of over 25 years. An inventory of 442 species is included, covering a wide range of
habitats: bare sand, seagrass beds, coral reefs, channels between the keys, and deep waters
over 600 feet in depth a few miles southward. Over 300 species were associated with coral
reef habitat. This study represents the first fish survey conducted on the Florida Reef Tract.
Field observations were made largely with the use of a diving helmet, which enabled the
investigator to observe and photograph the fish in their native habitat, and to give
information as to their behavior, feeding and habits, and especially adaptive coloration.
Much of Longley's work is documented by the world's first underwater color photography.
Following the death of Dr. Longley, Dr. Hildebrand undertook the editing and the
completion of the manuscript. Material added by him bear his initials.
213. Lucké, B.. 1937-1938. Studies on the tumors of fishes (of the snapper family Lutjanidae) . Carnegie
Institution of Washington, Year Book.
Note: published as follows; 1937, v.36, p. 98-99; 1938, v.37, p. 92-94.
Certain kinds of tumors found on thirty nine fish belonging to several species of snappers
closely resemble human neoplasms arising from nerves. They arise in the subcutaneous
tissue and appear as flattened oval masses. No tumors of this kind were found on other
species of fish. Epithelial growths were found in thirty specimens of Halichores radiatus,
from a total of six thousand fish observations. Multiple papillomas of the skin and the eye
were reported in a green turtle caught off Cape Sable.
214. ————. 1942. Tumors of the nerve sheaths in fish of the snapper family (Lutjanidae). Archives of
Pathology 34: 133-50.
Fish of the snapper family Lutjanidae are commonly afflicted with tumors which resemble
the nerve sheath tumors of man called variously neurinoma, neurolemmoma, schwannoma,
or neurofibroma. Neoplasms of this kind have been observed in 76 fish of three species,
the gray snapper (Lutjanus griseus), the dog snapper (L. jocu), and the schoolmaster (L.
apodus). Most of the fish were collected from the Dry Tortugas. Many other fish families
54
were examined, however no tumors of the kind were found. The tumors generally were
found along the course of the subcutaneous nerves, particularly of the head and dorsal
regions, as solitary or multiple, relatively large firm white masses. Like human neoplasms,
the tumors of fish are usually composed of two kinds of tissue: one compact and richly
fibrocellular; the other loose reticulated and poorly cellular. The component cells and
intercellular fibers of the tumors appear to be essentially the same, and arranged in similar
patterns, in fish and man. Unlike human tumors, the fish tumors, though well
circumscribed, are usually not encapsulated. Nerve sheath tumors appear to be more
common in certain fish species than in man. The frequency of occurrence of these tumors,
which can be maintained for long periods in marine aquariums, renders them favorable
material for studies of neoplasms.
215. Lucké, B. and H. G. Schlumberger. 1949. Neoplasia in cold-blooded vertebrates. Physiological
Reviews 29, no. 2: 91-126.
This review complements an earlier review to source material, abstracts of all the reports in
the literature dealing with tumors in fishes, reptiles, and amphibians. In regards to fishes of
the Tortugas, a review is presented on the senior author's work on tumors of nerve tissue,
as described in Lucke (1942, reference no. 214).
216. Lynts, G. W. 1968. Analysis of recent foraminiferal fauna from the Dry Tortugas, Florida. (abs.).
Geological Society of America Special Paper 101: 128-29.
This analysis of total foraminiferal population is based upon 16 samples collected in 1960
from the Dry Tortugas, Florida. Fourteen samples represent reoccupation of stations
sampled by Cushman (1922) in his original description of the foraminiferal fauna. Q-
modal factor-vector analysis indicates that the fauna is characterized by three assemblages
(factors) which account for 89 per cent of the total information (sum of squares of all
entries in data table). In general, these assemblages are characterized by a few dominant
species. One of the assemblages, Assemblage III, is directly related (r= +0.911) to depth
of water. The total population of the 14 samples representing reoccupation of Cushman's
stations were compared with the total population indicated by Cushman (1922).
Comparison was made using F-ratios and percentage of number of species occurring in
both samples (Sc). F-ratios estimate degree of variation between samples, whereas Sc
measures variation in species composition. F-ratios indicated that at seven of the 14
stations there were significant differences in total population between the 1919 and 1960
collections. Sc's ranged from 18.3 to 56.9 per cent and showed no relationship to degree of
variation estimated by F-ratios. This variation in foraminiferal fauna between collections
may represent either real changes in populations or apparent variations. If variations are
real, they reflect changes in the ecosystem over the 41 years. If variations are apparent,
they may represent either inability to resample microhabitats or vagaries in taxonomic
discrimination.
217. Lyons, W. G. 1980. Polyplacophora of Dry Tortugas Florida with comments on /schnochiton
hartmeyeri. Bulletin of the American Malacological Union, Inc. 46: 34-37.
450 specimens and 14 species of chitons were collected during 1978-79 from a station near
Garden Key. Only Acanthochitona sp. and Stenoplax purpurascens were relatively
common.
218. Mann, A. 1936. Diatoms in bottom deposits from the Bahamas and the Florida Keys. Papers From
the Tortugas Laboratory 29: 121-28.
Note: This is Appendix 1 to Calcareous shallow water marine deposits of Florida and the
Bahamas by Eldon Marion Thorp .
The twenty-four samples of calcareous sand collected by Doctor Vaughn in 1914 between
Cape Florida and Key West and at Tortugas were examined at the time these samples were
.
.
|
|
q
25)
received. A list of stations at which diatoms were collected is given, including Tortugas,
with a list of the diatoms found. Species of the genus Mastogloia are very abundant in
these Florida samples, and the author has found them to be so in all collections from
Florida Waters. In other parts of the world they are relatively much less abundant.
219. Manter, H. W. 1942. Gasterostomes (Trematoda) of Tortugas, Florida. Papers Tortugas Laboratory
33: 1-19 (issued June, 1940).
Carnegie Institution of Washington Publication Number 524.
A report of the Gasterostomatus trematodes collected in 1930, 1931, and 1932 is given
here. Fifteen species are reported; nine are considered new.
220. . 1934. The genus Helicometra and related trematodes from Tortugas, Florida. Papers
Tortugas Laboratory 28: 167-80 (issued Mar. 1933).
Carnegie Institution of Washington Publication Number 435.
Observations on the trematode genera Helicometra, Helicometrina and a new related form
are based on material collected at the Carnegie Biological Laboratory at Tortugas, Florida.
The genus Helicometra is represented at Tortugas by three species, H. execta, H. torta, and
H. fasiata. The characteristics of each of these are described. A key is given to the species
of the genus. H. execta is recorded from 6 additional hosts, making a total of 10. A
mutilated specimen of H. torta showed this species has little or no power of regeneration.
Helicometra fasciata from three hosts at 50 to 60 fathoms is reported for the first time from
America. Metacercariae of Helicometrina nimia are described encysted in the muscles of
the shrimps, Lysmata intermedia and Crangon formosum. Cercariae from Columbella
mercatoria identified as Cercaria J of Miller were found to encyst readily in the muscles of
Lysmata intermedia. Helicometrina parva, a new species is described .
221. ————. 1934. Some digenetic trematodes from deep-water fish of Tortugas, Florida. Papers
Tortugas Laboratory 28: 257-345 (issued Jan. 16, 1934).
Carnegie Institution of Washington Publication Number 435.
The parasitic fauna of ocean depths is practically unknown. Extensive fish population
occurs at all depths, very little study has been made on the helminths of these fish. During
the summers of 1930, 1931, and 1932 collections were made of parasites from fish trawled
from depths varying from 40 to 582 fathoms at Tortugas, Florida. Most of these hauls
were made about 10 miles south of Loggerhead Key. Fish taken from these depths were
commonly parasitized by helminths and especially by trematodes. It was found that the
trematode fauna of the deep-water fish is practically as abundant and as varied as is the rich
trematode fauna of the reef fish. A description is given of 49 species of trematodes
collected from approximately 90 species of fish from depths of 40 to 582 fathoms. 721
individual fish were examined. Approximately 80 per cent of the host species were
infected with trematodes, a percentage comparable with the degree of infection found in
fish of shallow water. One new subfamily (of the family Heterophyidae), 11 new genera
and 33 new species are described. Seven species of trematodes, from deep water only at
Tortugas, are identical with forms well known from northern regions. Studies from shallow
water at Tortugas show practically no similarity to northern forms. The deep-water
trematode fauna is more like the surface fauna of Maine, Great Britain or Norway than like
the shallow-water fauna at Tortugas, only a few miles away. This tendency to resemble
surface trematodes of cold-water regions suggests that temperature is an important factor in
the distribution of marine fish trematodes. This study emphasized the fact that a gradient of
changing environment (such as depth) is reflected not only in the free-living population of
a region but also in their parasites.
222. Marsh, G. 1940. The effect of light on the inherent E. M. F. of Valonia ventricosa. I. Intensity and
time relations. Papers Tortugas Laboratory 32: 65-84 (issued Oct. 1939).
56
Carnegie Institution of Washington Publication Number 517.
The interpretation of the electrical changes produced in green plants by light has been
retarded by the confused nature of the published results. In order to interpret electrical
changes in a tissue in terms of some underlying process it is essential to obtain precise
information concerning the distribution of E.M.F. within the tissue and the conditions of
summation of the potentials of the individual cells included in the electrical circuit. The
present paper reports the effect of visible light at known intensities upon the inherent
E.M.F. of the coenocytic alga Valonia ventricosa. Results indicated that when the intensity
of incident light is altered, the inherent E.M.F. of an impaled Valonia cell undergoes a
characteristic cycle of change with definite time relations, following which a steady level is
reached. The steady level of E.M.F, plotted against the logarithm of the light intensity rises
from the dark potential along a sigmoid curve to a maximum at about 250 foot-candles,
then descends along a similar curve toward the dark potential. The decline in potential with
light intensity beyond the maximum was reversible. No injury was detected at any
intensity. The effect of intermittent light with equal light-dark periods was similar to that
of continuous light of half the intensity, save in one experiment, wherein the effect was
similar to that of continuous light of increased intensity. It is concluded that the effect of
light on the E.M.F. is due primarily to the release of oxygen in photosynthesis.
223. ————. 1940. The effect of light on the inherent E. M. F. of Valonia ventricosa. II. The relative
energy absorption spectrum. Papers Tortugas Laboratory 32: 99-120 (issued May 1940).
Carnegie Institution of Washington Publication Number 517. :
The interest in the relative effectiveness of different wave-length bands of visible light
upon bioelectric potentials centers about the question of the nature of the agent in the living
cell which absorbs the radiant energy, and its contribution to the electromotive mechanism.
For the green plants three principal lines of evidence have been adduced to support the
conclusion that chlorophyll is the photosensitive agent. Results indicated that the steady
E.M.F. of impaled Valonia ventricosa illuminated with light of limited spectral
composition was matched with white light. The ratio of the intensity of white to that of
filtered light for an E.M.F. match was independent of the magnitude of the E.M.F. matched
and of the absolute intensities. The relative energy absorption is shown to compare fairly
well for the filter series with the relative absorption of chlorophyll mixtures over the same
spectral range calculated from the determinations of the absorption coefficient published by
Zscheile. It is concluded that chlorophyll is the photosensitive material absorbing the
radiant energy responsible for the effect of light upon the protoplasmic E.M.F. in Valonia.
The chlorophyll system is, therefore, an intimate part of the electromotive mechanism.
224.
. 1937. Effect of temperature upon the inherent potential of Valonia. Papers Tortugas
Laboratory 31: 1-16.
Carnegie Institution of Washington Publication Number 475.
The effect of temperature upon the potential is of prime importance in the determination of
the nature of the underlying electrochemical process. The electromotive force of a system
in thermodynamic equilibrium (including the diffusion potential) is proportional to the
absolute temperature (Q 10 of 1.04 or less within the biological range of temperatures).
The E.M.F. found across the protoplasmic layer of Valonia is not a thermodynamic one. It
is produced by an oxidation-reduction system in flux equilibrium at phase boundaries
within the cell. The E.M.F. is not primarily determined by the external medium. The
influence of the salt content of the sea-water upon the inherent potential is fundamentally
no different from the influence of the composition of the medium upon any other biological
process, as respiration, irritability, contractility, etc., where specific electrolytes in different
proportions condition, but do not cause, the process.
S/
225. Mast, S.O. 1911. Behavior of the loggerhead turtle in depositing its eggs. Papers Tortugas
Laboratory 3: 63-67.
Carnegie Institution of Washington Publication Number 132.
The nesting behavior of a single loggerhead turtle is described.
226. Matthai, G. 1915. Preliminary report on the comparative morphology of the recent Madreporaria
around Tortugas. Carnegie Institution of Washington, Year Book 14: 209.
General observations were recorded on the common corals of the Tortugas. The only
species that extruded larvae was Favia fragum.
227. Mayer, A. G.. 1908. The annual breeding swarm of the Atlantic Palolo. Papers Tortugas
Laboratory 1: 105-12.
Carnegie Institution of Washington Publication Number 102.
The habits of the "Atlantic palolo" are quite similar to those of the palolo worm of Samoa
and the Fiji Islands. The worms are, however, specifically different, the Atlantic palolo
being Eunice fucata Ehlers, and the Pacific worm E. viridis Gray. The annual swarming of
the Atlantic palolo has been observed only at Tortugas, Florida, although the worm is
abundant in the Bahamas and other parts of the West Indies.
228. . 1911. The converse relation between ciliary and neuro-muscular movements. Papers
Tortugas Laboratory 3: 1-25.
Carnegie Institution of Washington Publication Number 132.
A series of experiments on marine invertebrates were conducted beginning at the Tortugas
Laboratory, and later at Woods Hole, Mass, and the New York Aquarium involving the
effects of ions of blood salts, magnesium, sodium, calcium, ammonium, potassium, and
hydrogen on neuro-muscular systems in relation to maintaining ciliary movements. In each
case they are the exact apposite of their effects upon ciliary movements of invertebrates
studied. Studies were carried out on invertebrate organisms abundant at Tortugas
including annelid larvae, Limulus, veligers , actinian larvae, larvae of the Atlantic palolo
worm, Eunice fucata, and ctenophores, Cassiopea. Preliminary reports of the research
were published in the Biological Bull., Woods Hole, v. 17 (341-342); in the Proceedings
of the Soc. for Experimental Biology and Medicine, 1909, No. 7, (19-20), and in the
Carnegie Year Book for 1909, p. 152.
229. ————. 1914. The effects of temperature upon tropical marine animals. Papers Tortugas
Laboratory 6: 1-24.
Carnegie Institution of Washington Publication Number 183.
Tropical marine animals commonly live within 5° C. of their temperature of maximum
activity and within 10° to 15° C. of their upper death temperature. In marine tropical forms
even a few degrees of heat or cold cause a marked depression in movement. In tropical
Scyphomedusae this depression of movement appears to augment about as the square of
the change in temperature from that of the optimum. Time is an important factor in these
experiments, for animals can withstand a higher degree of heat if the temperature be raised
quickly than if it be raised slowly. It appears that the reef corals at Tortugas, Florida, live
in water which is commonly within 10° C. of their upper death-temperature, and if the
ocean were heated to 38° C. (100.4° F.) only one species, Siderastraea radians, could
survive. Next to Siderastraea radians the most resistant coral is S. siderea. It is associated
in its habitat with Orbicella annularis one of the most sensitive of the reef corals, which is
killed at 14.1° and 36.8° C. In general, however, the corals of the shallow-reef flats, such
as Siderastraea radians, Porites furcata, and Maeandra areolata, are the most resistant
both to heat and cold, while those of deep water, such as Madrepora palmata , Eusimilia
knorri, and Oculina diffusa, are the least resistant. As a result, we are led to conclude that
were the water cooled by an exceptionally prolonged norther to 13.9° C. for 9 hours,
58
230.
23s
Siderastraea radians , S. siderea, and Maeandra areolata would survive without apparent
injury while Porites furcata, P. clavaria, Maeandra clivosa, and Favia fragum would also
survive, but with more or less injury. This temperature would be fatal to Orbicella
annularis, Porites astraeoides, and Madrepora muricata (cervicornis).
. 1922. Hydrogen-ion concentration and electrical conductivity of the surface water of the
Atlantic and Pacific. Papers Tortugas Laboratory 18: 61-85.
Carnegie Institution of Washington Publication Number 312.
The hydrogen-ion concentration of sea-water was determined by placing 0.4 c.c. of 0.1 per
cent of the red dye thymolsuphonemphthalein in 70 per cent alcohol, in a test-tube of
resistance glass, 24 mm. in caliber, then adding sea-water so as to make up 30 c.c. of
solution. A series of such tubes, ranging from 7.95 to 8.3 pH, was standardized by
Professor J.F. McClendon and presented to the author who restandardized these tubes at
intervals of two years by comparison with determinations of pH made by a Leeds and
Northrup potentiometer. In order to avoid writing negative exponents, the symbol "pH" to
indicate the negative logarithm of the hydrogen-ion concentration was devised. Despite its
artificiality, one soon finds that the pH system gives a clearer idea of the alkalinity or
acidity of a solution than does a direct expression of the hydrogen-ion concentration. In
testing water, pH 7 would indicate practical neutrality; pH above 7, alkalinity; and below 7
acidity. The carbon-dioxide tension of the sea-water was calculated from the pH and the
temperature by the method devised by McClendon, Gault, and Mulholland (1917, Carnegie
Inst. Wash. Pub. No. 251, p. 36). McClendon found that the pH of sea-water falls 0.01 for
1° C. decline in temperature. The salinity of the sea is expressed in grams of total salts per
1,000 grams of sea-water, and was determined by the well-known method of using a
standard AgNO; solution with K,CrO, as an indicator, and testing against a sample of
standard sea-water obtained from Professor Martin Knudsen. Upon being taken from the
sea, the water was tested for temperature and pH, and a sample was preserved for
determination of salinity. In connection with these tests of hydrogen-ion concentration, the
electrical conductivity of the sea-water off Tutuila, Samoa and Tortugas, Florida was
determined by Kohlrausch's method. At Tortugas, Florida, the conductivity of sea-water
having 20.06 grams of chlorine in 1,000 grams of water, corresponding to a salinity of
36.24, was determined by the same apparatus, and with a portion of the same KCI solution
used in Samoa. In lagoons such as that of Tortugas, Florida, and in closed shallow areas,
McClendon found there was a diurnal variation in the pH, the water becoming more
alkaline by day and relatively acid during the night. This was attributed to the effect of
photosynthesis by plant life, which is active in daylight but ceases during the night. Over
shallow regions, where the water may become impounded in tide-pools at low tide, the
effect of photosynthesis is often very marked, the pH changing greatly while the
temperature may change but little. The rise in pH was due to the loss of Co) resulting from
photosynthesis.
. 1914. The law governing the loss of weight in starving Cassiopea. Papers Tortugas
Laboratory 6: 55-82.
Carnegie Institution of Washington Publication Number 183.
The medusae were always starved in the purest sea-water which was either dipped from the
ocean in glass or canvas buckets or pumped into glass reservoir tanks through hard-rubber
pipes by means of a hard-rubber pump. The medusae were starved side by side in one and
the same glass aquarium, but when this was impossible the aquaria were of similar size and
form and were placed side by side, so as to be subjected to similar environmental changes.
The decline in weight of two normal medusae of Cassiopea xamachana starved each in one
liter of sea-water, changed once in 24 hours, and kept in the diffuse daylight of the
laboratory at Tortugas, Florida, from June 8 to 20, 1913. One medusa, A, was starved in
sea-water which had been passed through two glass funnels each holding two sheets of
232%
233%
234.
235.
236.
ZBI
59
Chardin filter paper. The other medusa, B, was starved in sea-water, which, in addition to
having been filtered through the Chardin Filters, was also filtered through a bacteria-proof
porcelain filter. It appears that all food had been removed from the water by Chardin
filters and the medusa in the bacteria-free sea-water starved more slowly than the one in the
sea-water which had not been passed through the porcelain filter.
. 1908. Marine laboratories, and our Atlantic coast. The American Naturalist 42: 533-36.
In this general article by Dr. Mayer concerning the importance of marine laboratories along
our Atlantic Coast, the Tortugas is mentioned as having a certain remoteness from the
busy world and consequent freedom from interruption peculiarly favorable to the conduct
of research.
. 1915. The nature of nerve-conduction in Cassiopea. Proceedings of the National Academy
of Science 1: 270-274.
Narrative same as in reference no. 234.
. 1917. Nerve-conduction in Cassiopea xamachana. Papers Tortugas Laboratory 11: 1-20.
Carnegie Institution of Washington Publication Number 251.
By means of Professor E.G. Conklin and the authorities at Princeton the author used the
facilities at the Biological Laboratory in Guyot Hall, where the kymograph records taken at
Tortugas were studied and the results tabulated. The object of this research was to obtain
an accurate quantitative determination of the rate of nerve-conduction in natural and in
diluted sea-water at constant temperature, and also to estimate the effects of various
artificial sea-water solutions containing all or some of the sodium, magnesium, calcium,
and potassium cations of sea-water. The effects of temperature upon nerve-conduction are
also of great importance. These studies were carried out in June and July 1916, upon
Cassiopea xamachana, a thizostomous scyphomedusa which is abundant in the salt-water
moat surrounding Fort Jefferson at Tortugas, Florida. In summary, nerve-conduction is due
to a chemical reaction involving the cations of sodium, calcium, and potassium.
Magnesium is non-essential. Observations do not support the "local action" theory of Lillie
(1916).
. 1918. Nerve-conduction in diluted and concentrated sea-water. Papers Tortugas
Laboratory 12: 179-83.
Carnegie Institution of Washington Publication Number 252.
Ring-shaped strips of subumbrella tissue of the scyphomedusa Cassiopea xamachana were
deprived of marginal sense-organs and placed in concentrated sea-water in order to
determine the effect of concentration of electrolytes upon their rate of nerve-conduction.
Experiments made in 1917 upon Cassiopea found that the rate has increased while the
electrical conductivity has diminished. The injurious effects of concentrated sea-water
upon regeneration and growth have been studied by Loeb, and by Goldfarb (1914), and
there is a general resemblance between their curves and those in this paper for the rate of
nerve-conduction, excepting that for regeneration somewhat dilute sea-water seems to be
more favorable than normal sea-water, whereas in nerve-conduction the highest rate is
obtained in slightly concentrated sea-water .
. 1908. A plan for increasing the efficiency of marine expeditions, marine laboratories and
our Atlantic coast. American Naturalist 42: 533.
Narrative same as in reference no. 232.
. 1914. The relation between the degree of concentration of electrolytes of sea water and the
rate of nerve conduction in Cassiopea. Papers Tortugas Laboratory 6: 25-54.
Carnegie Institution of Washington Publication Number 183.
60
238.
ese).
240.
241.
If sea-water be diluted with distilled water, or with a 0.9 molecular solution of dextrose,
thus preserving its normal osmotic pressure but reducing the concentration of the cations of
sodium, magnesium, calcium, and potassium, the rate of nerve-conduction increases as
dilution proceeds, becoming most rapid in 90 per cent sea-water + 10 per cent distilled
water or dextrose. The sodium cation is an active stimulant for nerve-conduction.
Experiments with the magnesium cation show that it is not a stimulant for nerve-
conduction. In very slight excess the potassium cation produces a permanently stimulating
effect, as does sodium, but in denser concentration it produces momentary stimulation of
the rate of nerve-conduction followed by depression. In all essential respects the effects of
potassium are similar in kind, but more marked in degree, to those of sodium.
—. 1908. Rhythmical pulsation in Scyphomedusae.II. Papers Tortugas Laboratory 1: 113-31.
Carnegie Institution of Washington Publication Number 102.
The following paper presents the results of a continuation of studies, the first report of
which appeared in publication No. 47 of the Carnegie Institution of Washington, 1906.
The present paper aims to correct certain errors in the previous report, and to announce
some new results. Conclusions presented suggested that sea-water is a balanced fluid
neither inhibiting nor stimulating pulsation in Cassiopea xamachana. The stimulus which
causes pulsation is due to the constant formation of sodium oxalate in the terminal
entodermal cells of the marginal sense organs. This sodium oxalate precipitates calcium,
as calcium oxalate, thus setting free sodium chloride and sulphate which act as nervous
stimulants. Pulsation is thus caused by the constant maintenance at the nervous centers in
the sense-organs of a slight excess of sodium over and above that found in the surrounding
sea-water. In Cassiopea the pulsation-stimulus is conducted by the diffuse nervous network
of the subumbrella, and is independent of the muscles which may or may not respond to its
presence by contraction. Strong primary nervous and muscular excitement followed by
exhaustion and sustained muscular tetanus is produced in Lepas or in Cassiopea by a
solution containing the amounts and proportions of NaCl+KCl+CaCl? found in sea-water.
—. 1900. Some medusae from the Tortugas, Florida. Bulletin of the Museum of Comparative
Zoology at Harvard College 37, no. 2: 1-82.
This extensive article on the medusae of the Dry Tortugas includes an alphabetical list of
species described, table showing the wide geographical range on some Tortugas Medusae,
and comparisons of the Tortugas Fauna with that of Southern New England, the Tropical
Atlantic, Fiji Islands and Tropical Pacific.
. 1902. The Tortugas, Florida as a station for research in biology. Science (Washington,
DIE) Mg 190-192.
The advantages of establishing a tropical research marine biological laboratory in the Dry
Tortugas over other Caribbean sites are discussed. The nearness of the Florida Current, an
extension of the Gulf Stream, to the Tortugas is a distinct advantage over other sites
because of its richness in pelagics, especially small juveniles and larvae during the summer.
Other sites are richer in coral, such as Jamaica, but they are further from the Gulf Stream
and are impacted by land runoff.
. 1918. Toxic effects due to high temperature. Papers Tortugas Laboratory 12: 173-78.
Carnegie Institution of Washington Publication Number 252.
The experiments cited below appear to indicate that death from high temperature may be
due to the accumulation of acid in the tissues. Reef corals from Tortugas, Florida, were
kept at a constant temperature in warm ocean-water for 60 minutes in a thermostat, in the
dark the temperature remaining constant within about 0.1° C. throughout the hour. In this
manner the temperature was found that is just sufficient to kill the coral. The results are as
follows: Acropora muricata 34.7, Orbicella annularis 35.6, Porites astraeoides 35.8,
61
Porites clavaria 36.4, Maeandra areolata 36.8, Porites furcata 36.85, Favia fragum
37.05, Siderastrea radians 38.2. It is apparent that those corals which live in cool,
relatively agitated water, free from silt, are those that can not withstand high temperatures,
whereas those which live in the hot, silt-laden shallows near shore are, generally speaking,
forms which can resist high temperature. Favia fragum is however an exception. It seems
possible that death from high temperature may be due to the accumulation of acid (possibly
H,COs;) in the tissues, the rate of formation of this acid being related to the rate of
metabolism of the tissues. Thus animals of the same class having a high rate of
metabolism, as measured by oxygen consumption, are more sensitive to heat and to CO)
than those having a low rate of metabolism.
242. . 1922. The tracking instinct in a Tortugas ant. Papers Tortugas Laboratory 18: 101-7.
Carnegie Institution of Washington Publication Number 312.
Monomorium destructor Jerdon, a tropicopolitan ant of East Indian origin, was identified
in Florida . It is a small, reddish-brown ant, a great pest in the wooden buildings of the
Tortugas laboratory. These pests have killed rats confined in cages within 24 hours. The
experiments described were made on the flat wooden floor of the laboratory. To attract the
ants, a number of recently killed houseflies were impaled upon a pin. The pin with its lure
of flies was then thrust into the floor in front of a foraging ant, which would often pass
within 0.25 inch of the lure without perceiving the flies; but if its course were such that it
came appreciably nearer than 0.25 inch, the ant suddenly turned toward the flies, and
without apparent excitement appeared to "inspect" them, spending a half minute or more
crawling over them and stroking them with its antennae. This "finder ant" soon leaves the
flies without carrying off any piece of them, but instead of moving off in the erratic and
tortuous path it was pursuing before it found the flies, it now goes in a fairly straight path
toward some crevice in the floor, out of which there soon pours an excited swarm of its
nest-mates, who proceed toward the flies in a fairly straight path. When an ant returns to
the nest it pursues a fairly straight path which is more or less right in direction, but when
the ant has gone the correct distance, it begins to wander in more or less tortuous courses
until it finds the nest.
243. McClendon, J. F. 1917. Diurnal changes in the sea at Tortugas, Florida. Proceedings of the National
Academy of Science 3: 692.
The only diurnal change noted in the Gulf Stream was a change in temperature of about 1
degree C and the resulting change in oxygen tension. However, marked differences were
found in temperature, pH, CO >, and O; concentration in shallow water where light could
reach the bottom. The temperature, O, concentration and O, tension were lowest and the
CO), concentration and CO, tension highest at5 A.M. The temperature, O2 concentration
and O, tension were highest and CO, concentration and CO) tension were lowest at 3 P.M.
during July at Dry Tortugas. The magnitude and exact time of minima and maxima varied
from day to day, and varied a great deal with station location. Studies on the effects of
these changes on organisms were made. The limiting factor for plants was nitrogen, while
the limiting factor for animals was food and oxygen.
244. ———. 1917. Effect of oxygen tension on the metabolism of Cassiopea. Proceedings of the
National Academy of Science 3: 715-16.
Experiments on the effect of oxygen tension on the metabolism of Cassiopea were carried
out at the Tortugas Laboratory by using the umbrella of Cassiopea to maintain a layer of
cells in seawater at 30 degrees C. The metabolism varied with oxygen concentration. This
may be true for all animals, however there is a distinction between the metabolism of
vertebrate cells and Cassiopea. Vertebrate cells give out lactic acid when asphyxiated
whereas Cassiopea may remain without oxygen for seven hours without giving out CO, or
any other acid. Although details of the experiments will be published elsewhere, it was
62
245.
246.
247.
248.
concluded that changes in the threshold of stimulation of the respiratory and basomotor
centers may affect metabolism in man and animals.
. 1917. The effect of stretching on the rate of conduction in the neuro-muscular network in
Cassiopea. Proceedings of the National Academy of Science 3: 703.
The experiments on Cassiopea collected at the Dry Tortugas, tend to support the
conclusions reached by Carlson, that stretching the nerve does not change the rate of the
nerve impulse, and that the conducting substance itself, can be stretched and relaxed.
. 1917. The equilibrium of Tortugas sea water with calcite and aragonite. Proceedings of the
National Academy of Science 3: 612-19.
This report provides information on the continuing controversy on the solubility of
Calcium chloride in sea water. The precipitation of CaCO; at Tortugas was studied by
T.W. Vaughan, R.B. Doyle, and G.H. Drew. Drew observed that denitrifying bacteria,
Pseudomonas calcis obtained from sea water was capable of changing calcium nitrate to
calcium carbonate in culture media, and supposes that a similar process occurs in seawater.
This study attempts to determine the nitrates or nitrites. If the pH is maintained (by plants)
at 8.2 at the Tortugas, the introduction of calcite crystals would result in a lowering of the
calcium content of Tortugas seawater by about 4.5%.
. 1914. Experiments on the permeability of cells. Papers Tortugas Laboratory 6: 123-30.
Carnegie Institution of Washington Publication Number 183.
One of the most important steps in the analysis of life was the discovery of oxygen. Ever
since that time it has been known that animals absorb free oxygen and give it out in a
combined form. In this experiment three methods of procedure were followed: (1) the use
of cell masses as partitions (on eggs of Lytechinus); (2) the use of quantities of eggs
suspended in a liquid medium (on eggs of Fundulus); (3) experiments on individual eggs
(of Arbacia). The permeability of the egg to ions and perhaps some other substances
increases on fertilization. The unfertilized egg is perhaps in a dormant condition and the
increase in permeability probably allows a rapid interchange with the surrounding medium
necessary for activity (development). Whereas this supposed significance of permeability
has not been proven, the sea-urchin;s egg is not an exception. The relation of permeability
to oxidation can hardly be determined until more is known about the mechanism of animal
oxidations. These seem to depend on structure since complete oxidations cease when
structure is completely destroyed. Reference is made only to oxidations resulting in the
formation of CO:. Oxidizing enzymes such as tyrosinase, which are independent of
structure, do not completely oxidize the substances acted on.
. 1911. On adaptations in structure and habits of some marine animals of Tortugas, Florida.
Papers Tortugas Laboratory 3: 55-62.
Carnegie Institution of Washington Publication Number 132.
This article discusses the habits of some marine animals of the Tortugas. Many of these
animals were thigmotactic and remained in glass tubes rather than in the open. They
learned to find the tubes when removed from them. Such was the case with five species of
the Alpheidae, one of the Pontoniidae, Typion tortugae Rathbun, and Gonodactylus
aertedii. All the anemones were thigmotactic on their bases. These same animals were
heliotropic. The crustaceans were negatively heliotropic and the anemones kept their bases
from the light, while Cradactis variabilis Hargitt hid all but the tips of the fronds and
tentacles from the light. In removing its base from the light, Stoichactis helianthus, which
lives on coral heads, makes snail-like movements similar to Metridium, while Cradactis,
which lives in holes in decayed coral heads, crawls on its tentacles.
63
249. . 1918. On changes in the sea and their relation to organisms. Papers Tortugas Laboratory
12: 213-58.
Carnegie Institution of Washington Publication Number 252.
The sea and air form the circulating media for the living organisms of the world. The local
composition of the sea is distinctly affected by living organisms. The local changes in the
composition of the sea are the subject of the present paper. These changes are due chiefly
to organisms, but partly to meteorological causes. The water evaporated is returned with
addition of fixed nitrogen from electric discharges or falls on the land and is returned with
various salts, chiefly CaCO3, and with fixed nitrogen and other products of organisms.
Various seaweeds absorb CO) thus leaving an excess of CaCO, which has a very low
solubility and is constantly being precipitated in certain warm seas, and is precipitated
within the bodies of organisms in the surface waters of all seas. In working out the relation
of H-ion concentration (pH) to the solubility of CaCO; in sea-water, it was found that all
sea-water is supersaturated with CaCO3, and will lose some of it if shaken with calcite or
aragonite crystals. The pH is influenced by plant and animal life and arises at Tortugas to
8.35 during the day over well-lighted bottoms rich in vegetation, and falls to 8.18 during
the night. It may be said, therefore, that conditions in shallow water over eelgrass or other
seaweed or corals (with symbiotic algae) favor the precipitation of CaCO3. The question
arises whether the occasional high pH of Tortugas sea-water is sufficient to explain the
precipitation of CaCO3. The author's experiment showed that if the pH of sea-water should
be maintained (by the action of plants) at 8.2 while it was agitated with calcite crystals, the
loss of CaCO; would be about 0.001 N, or 0.0005 M, or 0.1 gram per liter. This would
cause a deposit of 10 kg. per square meter of bottom in water 100 meters deep. This would
cause a lowering of the total calcium content of Tortugas sea-water by about 4.5 per cent.
The actual precipitation of CaCO; was most noticeable in the Marquesas lagoon. At 4
p.m., July 30, the pH was 8.46 and there was a precipitate of CaCO; coming down in the
water and encrusting the eel-grass.
250. ————. 1917. The standardization of a new calorimetric method for the determination of the
hydrogen-ion concentration, CO, tension and CO, and O; content of sea water, of animal
heat, and of CO) of the air, with a summary of similar data on bicarbonate solutions in
general. Journal of Biological Chemistry 30: 265-88.
Experiments were conducted on pH, CO, tension, CO, and oxygen content of Dry
Tortugas seawater and seawater from other oceanic areas using a Leeds & Northrup
potentimeter and a 0.1 KCL calomel electrode. It was found that neither the salinity nor
the alkaline reserve in seawater of the tropical or temperate oceans change sufficiently to
noticeably change the relation of pH to CO; tension, although the alkaline reserve does
change sufficiently to affect the total CO) greatly.
251. McClendon, J. F., C. C. Gault and S. Mulholiand. 1917. The hydrogen-ion concentration, CO2
tension, and CO2 content of sea water. Papers Tortugas Laboratory 11: 21-69.
Carnegie Institution of Washington Publication Number 251.
Narrative same as in reference no. 250.
252. Meeder, J. F. 1979. Corals and coral reefs of the Dry Tortugas, Florida. in Guide to sedimentation
for the Dry Tortugas. R.B. Halley (Compiler), 46-47. S.E. Geological Society Pub.
This paper presents a description of the corals from two localities in the Dry Tortugas, on a
back reef environment of a fringing reef near Garden Key, and the second, a series of patch
reefs off Loggerhead Key. The general setting, ecology, distribution, and types of corals
are discussed for each locality. Forty-one of the forty-two species of corals reported at the
Tortugas are covered in a field key and later described in this paper.
64
253. Miller, H. M. Jr. 1926-1929. Behavior of trematode larvae. Carnegie Institution of Washington,
Year Book.
Note: Published as follows: 1926, v.25, p. 243-244: 1929, v. 28, p. 295.
Anatomical/morphological descriptions of six larval tremeatodes infesting the mollusk,
Cerithium litteratum, taken from Bird Key Reef Porites beds, were provided. Percentages
of occurrence of the 6 cercariae were given. The behavior of the members of this
morphological group are described in detail, including aspects of their life history.
254. Miller, R. A. and H. B. Smith. 1931. Observations on the formation of the egg of Echinometra
lacunter. Papers Tortugas Laboratory 27: 47-52.
Carnegie Institution of Washington Publication Number 413.
The study of the ovaries of Echinometra was undertaken in the hope that by the use of
some of the newer cytological methods, it might be possible to extend our knowledge of
the processes of odgenesis, particularly of those concerned in the formation of yolk in the
chinoid egg. This paper presents the observations that have been made, and the
conclusions that have been drawn. It was found that in Echinometra lancunter,
undifferentiated cells along the wall of the ovary are uniform in appearance, although they
are destined to develop into two entirely different kinds of cells. As development
proceeds, some of the cells become odgonia, while others enlarge and disintegrate to form
deutoplasmic bodies. These are more coarsely granular than was the cytoplasm from
which they were formed. The granules exhibit different affinities for stains. Fully formed
nutritive spheres are of two kinds, granular and non-granular. The former are composed
entirely of cytoplasmic, or of both cytoplastic and nuclear material. The nutritive spheres
group themselves around the o6gonia in a follicular arrangement. Eventually the nutritive
spheres enter the egg and disappear as such, forming the yolk content of the cytoplasm,
which becomes homogeneous and evenly granular. The deutoplasmic granules are smaller
and more diffuse in the mature eggs than in the odcytes. The nutritive spheres have been
shown to be composed of phospholipins suspended in a homogeneous medium. It is
probable that they are largely lecithin in content. Their origin is not known.
255. Millspaugh, C. F. 1907. Flora of the sand keys of Florida. Columbian Museum 118, no. (Bot. Ser.
2): 191-245.
A list of species and details of vegetation, as well as elevational descriptions and size
dimensions of sand keys westward of Key West, including the Tortugan group, were
compiled by the author during the winter through spring of 1904.
256. Mitchell-Tapping, H. J. 1981. Particle breakdown of recent carbonate sediment in coral reefs.
Florida Scientist 44, no. 1: 21-29.
Skeletal particles of the major components of the carbonate sediment of the reef shoal
environment were examined using the scanning electron microscope. This examination
revealed no set pattern of skeletal breakdown according to microarchitectural structure, as
postulated by the Sorby principle, but that such a breakdown depends on mineralogical
composition, wall thickness, grain size and pattern, density, and the amount of cementation
and bloerosion. To investigate general particle-size abundances and deficiencies in
carbonate sediment, samples were taken from the reef crest, back-reef-rubble and open-
sand ecozones of the reef shoal environments of sites from the Bahamas, Dry Tortugas,
Lower Florida Keys, Grand Cayman Island and the U.S. Virgin Islands. Size analyses of
these samples showed that the sediment is moderately sell-sorted, coarsely-skewed and
leptokurtic. Although particle-size abundances (or modes) exist in each individual site,
there is no particular particle-size abundance that is common to all the sites. It is inferred
that the particle abundances (or modes) for each site are a product of the sorting potential
of the wave energy and that this sorting potential is the major control of the breakdown of
65
sand-sized skeletal particles rather than the microarchitectural structure as proposed by the
Sorby principle.
257. Moritz, C. E. 1936. Embryology of the sea-hare, Aplysia protea and of Crangon armillatus.
Carnegie Institution of Washington, Year Book 35: 90.
Observations from aquaria are recorded on the early development of Aplysia, from embryo
to 5 days beyond hatching. Adults were collected from the moat on Garden Key and Bird
Key Reef.
258. Multer, H. G. 1975. Field guide to some carbonate rock environments; Florida Keys and Western
Bahamas. Fairleigh Dickinson University 40: 175 pp.
This report presents the most recent compilation on Holocence sediments in the western
Bahamas and in the Florida Keys including the Dry Tortugas. Selected geologic literature
pertinent to local environments is noted within the text, with full bibliographic citations
following each subject area. Carbonate sand beach and beach rock environments from the
Florida Keys, Bimini, and Loggerhead Key are compared. In summary, Holocene
sediments of these areas today present a vast array of textures and constituent particles
characteristic of environments which have been subjected to fluctuating sea levels and
storm action. Such data may be used to interpret ancient environments.
259. . 1971. Holocene cementation of skeletal grains into beachrock, Dry Tortugas, Florida.
Carbonate Cement. O. P. Bircker. Baltimore, Maryland: Johns Hopkins Press.
A discussion is presented on the origin of beach rock at Loggerhead Key, Dry Tortugas.
Present evidence suggests that cementation is due to alternate wet and dry salt water spray
conditions with skeletal grains providing nuclei for precipitation from a supersaturated
calcium carbonate solution. The limited ground water conditions and lack of grain solution
for providing aragonite cement are two evidences in favor of the above cited evaporation
origin for the cement in this rock. Ginsburg (1953) reached similar conclusions for beach
rock in the same area.
260. Murphy, L. E. 1993. Dry Tortugas National Park, Submerged Cultural Resources Assessment, L. E.
Murphy. Submerged Cultural Resources Unit, Southwest Region, National Park Service,
Santa Fe, New Mexico.
This volume describes and assesses the known and potential archeological resources in
Fort Jefferson National Monument, later redesignated Dry Tortugas National Park. The
emphasis is on submerged cultural sites, particularly shipwrecks. The importance of
linking the natural resources with submerged cultural resources is provided by identifying
the biological influences on the cultural resources. The Dry Tortugas and South Florida
geological development and environmental succession is summarized with focus on the
postglacial development of the Florida-Reef Tract, depositional environments, coral reef
and sand key development as well as Late-glacial and Postglacial succession of
environments. An overview on the physical oceanography of the eastern Gulf of Mexico
concentrating on the Dry Tortugas with emphasis on currents and climate that affect
shipping vessel casualties and site preservation is provided. Recommendations for future
research and resources management are given.
261. Nance, J. M., E. F. Klima and F. J. Patella. 1986. Review of the Tortugas pink shrimp fishery from
May 1984 to December 1985, Galveston, Texas, NOAA/NMES, Southeast Fisheries
Center, Galveston, Texas. NOAA Tech. Memo.177.
Commercial pink shrimp fishing data from the Tortugas (Dry Tortugas Islands, Florida)
fishery were reviewed for biological year 1984 (May 1984-April 1985) and the first 8
months of biological year 1985 (May 1985-December 1985). Pink shrimp landings were
Just over 11.0 million pounds in biological year 1984 with 17,000 days of fishing
66
expended. This computed to a CPUE value of 643 pounds per day. Pink shrimp landings
for biological year 1985 are estimated to be around 9 million pounds with 15,000 days of
fishing expended. The predicted CPUE value for 1985 should be around 600 pounds per
day. Biological year 1984 experienced two extended periods of pink shrimp recruitment
into the Tortugas fishing grounds.
262. O'Neill, C. W. 1976. Sedimentology of East Key, Dry Tortugas (abs.). Florida Scientist 39 (Suppl.
263.
1), Fortieth Annual Meeting of the Florida Academy of Sciences at Eckerd College, St.
Petersburg, Florida March 18,19,20, 1975: 10.
East Key of the Dry Tortugas rests on a large crescent shaped bank and oscillates about a
stable core in response to seasonal variations. A simple strand/dune plant community is
largely responsible for short term stabilization of this central core. Historical studies
covering 200 years show that East Key varies in its bank position on long term basis and
has in the past been of much greater areal extent. In addition, historical records show the
Dry Tortugas group has decreased in extent from 11 keys to the present seven. This effect
is thought to be due to a combination of eustaic sea level change and storm degradation.
. 1976. "Sedimentology of East Key, Dry Tortugas, Florida." University of South Florida.
Ph.D. Dissertation.
The Dry Tortugas platform is a complex of reefs, banks, and shoals which lie 65 nautical
miles west of Key West, Florida. This reef platform most closely resembles the resorbed
reef of Maxwell's (1968) classification. A review of historical records covering a 463-year
period indicates that the Dry Tortugas island group is undergoing progressive degradation
and has been reduced from 11 rocky islets in 1513 to 7 at present. This reduction is
thought to be due to the combined effect of episodic events and the Holocene
transgression. East Key maintains a position on the windward segment of the Tortugas
group. Short-term changes to the key are basically in response to regular seasonal variation
and cause the island to oscillate to the north and south about a plant-stabilized core. Long-
term changes reflect the effect of episodic events and entail inundation, major shifts
inposition, and changes in the orientation of the key. Mean grain size of East Key
sediments is near 0.900 or coarse sand. The sediments are nearly without exception
moderately well sorted with an average value of 0.570. The mean skewness value is -0.22
(coarsely skewed). Texture was compared to morphology. The most striking correlation
occurred on the foreshore where there is a distinct tendency for sediments to become
coarser moving seaward from the berm. The average percentage composition of East Key
sediments, by constituent, was found to be 52 percent Halimeda, 35 percent coral, 6
percent Mollusca, 3 percent coralline algae, 1 percent Foraminifera, 1 percent echinoid
fragments and 2 percent miscellaneous plus unknown. The most significant correlation
between composition and morphology occurred on the foreshore, where percent Halimeda
generally increased seaward from the berm. This general increase in percent Halimeda
correlates with a tendency for sediments to become coarser moving seaward from the berm.
264. Ogden, J.C., W.C. Jaap, J. W. Porter, N. P. Smith, A. M. Szmant and D. Forcucci. 1993.
SEAKEYS: A large scale interdisciplinary study of the Florida Keys Reef Tract (abs.).
Proceedings of the Seventh International Coral Reef Symposium, v. 2. Mangilao, Guam:
University of Guam Marine Laboratory.
The SEAKEYS program has established a research framework which encompasses the
large geographic and long time scales of natural marine processes and ecosystem variation.
The core of the program is a series of instrumented, satellite-linked monitoring stations
which span the 220 mile coral reef tract. Mesoscale physical oceanographic studies are
concentrated in the major channels potentially linking Florida Bay and the population
centers of the Keys with the reef tract. Simultaneously, nutrient studies are probing the
possibility of sewage and agrichemical contamination, complicated by natural sources of
67
nutrients. A series of long-term photomosaic stations have tracked coral community
dynamics for more than 5 years. The design of the SEAKEYS program may provide an
example for long-term research on coral reefs elsewhere.
265. Osburn, R. C. 1914. The Bryozoa of the Tortugas Islands, Florida. Papers Tortugas Laboratory 5:
181-222.
Carnegie Institution of Washington Publication Number 182.
In the summer of 1908 the writer had the privilege of spending the month of June at the
Carnegie Institution Laboratory of Marine Biology on Loggerhead Key of the Tortugas
Islands. The entire period was devoted to a close search for the bryozoa inhabiting the
shallow waters about the reefs, on the piles of the old government dock on Garden key, in
the moat of old Fort Jefferson on the same key, and in dredging the shallow waters about
the islands down to 22 fathoms. Comparatively little work has been done on the bryozoa
of the Florida and West Indian regions. By comparing with lists from other regions where
the bryozoa have been carefully worked, it will be seen that the bryozoan fauna of the
Tortugas and of the Florida-West Indian regions 1s fairly rich in species and fairly
representative of tropical and semi-tropical regions.
266. Payne, F. 1937. Early development of Ptychodera bahamensis. Papers Tortugas Laboratory 31:
71-76 (issued July, 1936).
Carnegie Institution of Washington Publication Number 475.
The author records some observations made on Ptychodera bahamensis during the summer
of 1933 while working at the Carnegie Laboratory at Tortugas, Florida. The animals were
found most abundantly on the coral reef near Fort Jefferson in water which at low tide was
from one to two feet deep. The larvae described by Weldon, Morgan, and Stiansy and
assigned to Ptychodera bahmensis do not agree with the author's own observations of the
larvae of Ptychodera bahamensis which have been followed from the fertilized egg. It
seems conclusive that more than one species is involved or that errors have been made on
the part of someone. Even though the author has not followed development through to
metamorphosis, it seems clear that the tornariae, described by Weldon and Morgan are
different from the tornaria of Ptychodera bahamensis, assuming his identification is
correct.
267. . 1933-1938. Embryology and cytology of the balanoglossid, Ptychodera bahamensis...
Carnegie Institution of Washington, Year Book.
Note: published as follows: 1933, v. 32, p. 277-78; 1938, v. 37, p. 84.
Embryological work was conducted on the supposed protochordate, Ptychodera
bahamensis, starting with the fertilized egg. Development was followed as far as the
tornaria.
268. Pearse, A. S. 1934. Animals in brackish water ponds and pools at Dry Tortugas. Papers Dry
Tortugas Laboratory 28: 125-42 (issued Dec. 1932).
Carnegie Institution of Washington Publication Number 435.
During the summer of 1931 two ponds and three pools were studied on Long and Garden
Keys, Dry Tortugas. Because of the remoteness of the Dry Tortugas, the animals which
live in these isolated habitats are of particular interest. Results were as follows: Five
brackish water pools and ponds were studied on Long and Garden Keys. These were
variable in salinity and temperature, and limited in extent. They showed various stages of
evolution from sea to fresh water and conditions of life in them were more or less severe.
The animals in the ponds and pools were resistant to environmental variations. They lived
in salinities between 0.6 and 6.4 per cent and endured temperatures above 42° C. Some of
the small bodies of brackish water on Dry Tortugas contained curious mixtures of marine
and fresh-water animals. Callianassas, marine snails, mullets, and needle-fishes lived with
68
269.
270.
PHIM
Piles
dragon-fly nymphs, water boatmen, surface bugs, aquatic beetles and midges. Lists of the
animals collected and observed in each pond or pool are given. Each pond or pool
contained certain characteristic animals, which had become dominant in that particular
habitat. As the small pools trended toward fresh water, insects were increasingly dominant
in them. Insect populations show great pressure and spread into all available habitats.
When small pools are cut off from the ocean and are gradually transformed into fresh-water
habitats, insects are the pioneers and soon become dominant.
. 1934. Freezing points of bloods of certain littoral and estuarine animals. Papers Tortugas
Laboratory 28: 93-102 (issued Dec. 1932).
Carnegie Institution of Washington Publication Number 435.
It is well known that the bloods and body fluids of marine invertebrates and elasmobranchs
have about the same osmotic pressure as the ocean water in which they live, though the
salt content of such fluids is usually a little less than that of the surrounding medium.
Those of teleost fishes (marine, fresh-water and land) and of fresh-water and land animals
generally have osmotic pressures which are much below those of sea-water. Littoral
crustaceans and fishes are of particular interest because they represent various stages of
adjustment to life on land. At Tortugas, conditions are particularly favorable for the study
of such adjustments for the littoral group. The observations cited in this paper appear to
justify the conclusion that crabs and fishes which take up terrestrial life or air-breathing
have bloods of lower osmotic pressures than comparable marine or fresh-water animals.
The attainment of land life by marine animals is apparently associated with a reduction in
salinity and stabilization of the contents of the blood.
. 1934. Inhabitants of certain sponges at Dry Tortugas. Papers Tortugas Laboratory 28: 117-
24 (issued Dec. 1932).
Carnegie Institution of Washington Publication Number 435.
Many sponges are veritable living hotels. Their canals are densely populated with a variety
of animals and some species have not been found elsewhere. During the summer of 1931
the writer studied the animals which occurred in five species of sponges at Dry Tortugas.
The number of animals which live in big sponges is enormous. Though each sponge
appears to be occupied to its full capacity, the number of animals per cubic centimeter of
sponge is apparently influenced by depth and the size of the sponge itself. In loggerheads
there are relatively more guests in smaller sponges and in deep water. Among the species
of small sponges there are striking differences in the number of animals present and these
may be due to inherent qualities. For example Stematumenia foetida (Schmidt) contained
very few animals, whereas the slightly smaller Spongia officinalis L. was crowded.
. 1918. Notes on certain amphipods from the Gulf of Mexico, with descriptions of new
genera and new species. Proceedings of the U.S. National Museum 43, no. 1936: 369-79.
This is a report on a portion of the amphipods from the Gulf of Mexico in the collection of
the United States National Museum. The collections are from several sources and extend
over a long period of years. The greater part of them had not yet been examined. Those
described were taken chiefly by the steamers Fish Hawk and Albatross of the United States
Bureau of Fisheries.
. 1929. Observations on certain littoral and terrestrial animals at Tortugas, Florida, with
special reference to migrations from marine to terrestrial habitats. Papers Tortugas
Laboratory 26: 205-23.
Carnegie Institution of Washington Publication Number 391.
In the past, various types of animals have migrated from the ocean into fresh-water streams
or lakes, and from there gained a foothold on land. On the shores of all oceans, animals
may be found which are partially adjusted to life in fresh water or on land. The Dry
69
Tortugas contain no fresh water, and therefore offer an excellent opportunity to study
littoral animals which have no immediate contact with fresh-water habitats. In summary, at
Tortugas certain reef, beach and land animals were studied with reference to migrations
from sea to land. Hermit crabs which have become more or less adjusted to life on land
show a progressive reduction in the number of gills. Crabs which have migrated landward
show a progressive lessening of gill-volume. Beach animals which show any landward
trend usually live longer when kept in air than when kept in fresh water. Animals which
have attained some degree of ability to live on land have often also acquired a greater
degree of ability to resist the extraction of constituents of body fluids into fresh water.
Animals which migrate from the sea and become established on land do not do so on
account of one "lure" or one "danger." Each habitat has certain advantages and certain
disadvantages. A continually changing animal must continually make adjustments to a
continually changing environment, and when it migrates to a new habitat, must make many
compromises between new advantages and dangers, old necessities and new requirements,
and old habits and new abilities.
NB). . 1934. Observations on the parasites and commensals found associated with crustaceans and
fishes at the Dry Tortugas, Florida. Papers Tortugas Laboratory 28: 103-15 (issued Dec.
1932).
Carnegie Institution of Washington Publication Number 435.
During the summer of 1931 the writer had opportunity to conduct post mortems on various
crustaceans at Dry Tortugas from June 3 to August 22. Dr. Waldo Schmitt furnished and
identified many of the specimens. The following parasitic isopods were taken from fishes:
2 Cymothoa oesturm (L.) from the gill cavity of Caranx ruger (Bloch), July 26; 1
Rocinela signata Schioedte and Meinert from the gills of Promicrops itaiara Lichtenstein
in July and 1 from the gills of Lutianus analis (Cuvier et Valiencennes), June 27; 8
Excorallana tricornis (Hansen) from the nose of Promicrops itaiara Lichtenstein, July,
and 31 from the gill cavity of Epinephelus moro (C. et V.). The occurrence of the
parasites and commensals associated with crustaceans depends upon a variety of factors -
host specificity; habitat; habits, structure and physiology of hosts and parasites, etc. In
general the greatest number of species of parasites occurred in or near the littoral zone.
However, great numbers of parasites per host were encountered among some land
crustaceans.
274. ————. 1929. Two new mites from the gills of land crabs. Papers Tortugas Laboratory 26: 225-30.
Carnegie Institution of Washington Publication Number 391.
During July and August 1928, mites were found at Dry Tortugas on the gills of the land
hermit crab, Cenobita diogenes (Latreille), and on the Nassau crab, Gecarcinus lateralis
(Freminville). These crustaceans visit the ocean only once each year when they hatch out
their young. No mites were found on the gills of the ghost crab, Ocypoda albicans (Bosc),
which often visits the ocean and bathes its gills. Mixed or pure cultures of bacterial
populations are incapable of precipitating CaCO; in a sea-water medium if KNO3 and
organic matter as the sugars or similar forms free from calcium are added to the medium. A
number of different forms of bacteria in the sea possess the power of precipitating CaCO;
in appropriate media containing a large excess of soluble salts, but only under such
conditions. Among such organisms there is great variability to perform the task in
question, depending on the composition of the medium.
275. Perkins, H. F. 1908. Notes on the medusae of the Western Atlantic. Papers Tortugas Laboratory 1:
133-56.
Carnegie Institution of Washington Publication Number 102.
The Marine Biological Laboratory in the Dry Tortugas is well situated for the study of
many of the lower marine animals, their behavior, and the conditions of life, particularly
70
the coelenterates. One quite unique feature occurs in the Tortugas in the presence of the
old fortification and surrounding moat of Fort Jefferson. The moat affords remarkably
favorable conditions for the growth and multiplication of the lower forms of plants and
animals sheltered by the sea-wall, its shallow water warmed by the sun and kept from
stagnation by the agitation and partial change of the tides. The writer has for several years
been interested in the causes of migration and segregation of Medusae and has had the
privilege of examining specimens of this genus from Jamaica, and has studied the
characteristics of the specimens found in the Bahama Islands and at the Tortugas. There
was less of peculiarity in all the surroundings, the temperature of the water, storm
influence, and food supply being normal for the shores of coral islands. The only points of
difference to be noted in the medusae are with reference to size and color-pattern. The
main features of the two species, Cassiopea xamachana and Polyclonia frondosa are
presented.
276. Petrovic, C. A. and J. King Jr. 1973. Bird records from the Dry Tortugas. Florida Field Naturalist
277.
278. Phillips,
279.
1, no. 1: 5-8.
During a visit March 25 to April 4, 1967 to the Tortugas, a total of 70 species of mostly
land birds were recorded. Detailed observations on twenty specimens rarely seen at the
Tortugas are presented.
. 1972. Common elder and king rail from the Dry Tortugas Florida. Auk 89, no. 3: 660.
The authors watched the early spring migration at the Dry Tortugas Islands, which lie in
the Gulf of Mexico about 70 miles west of Key West, Florida, and recorded 70 species, the
majority land birds. The two records reported here represent significant additions to the
species known distribution .
A. H. 1917. Analytical search for metals in Tortugas marine organisms. Papers Tortugas
Laboratory 11: 89-93.
Carnegie Institution of Washington Publication Number 251.
This study concerns the problem stated in Year Book no.14, page 193 of the Carnegie
Institution of Washington. A large number of specimens were collected to be analyzed for
metals. The metals determined were iron, manganese, zinc, copper, and lead. For the
determination of zinc, copper, and lead, when the dried material was sufficient, 20 grams
were used as a sample; when it was not possible to use 20 grams, the results are all
calculated to 20 grams..
. 1922. Analytical search for metals in Tortugas marine organisms. Papers Tortugas
Laboratory 18: 95-99.
Carnegie Institution of Washington Publication Number 312.
Included in the material collected at the Tortugas and analyzed for metals (some of the
results of which were reported in the annual report of the Carnegie Institution for 1917)
was a brown spotted holothurian, Stickopus mobii, which was analyzed by the methods
there indicated. The element vanadium was found in the holothurian material and
heretofore has never been reported from seawater. Vanadium has been reported from
freshwater and in the blood of an acidian from the Bay of Naples. This vanadium content
of the blood does not seem to be a characteristic of all acidians, as two other species from
the Tortugas yielded no vanadium, neither did two other species of holothurians yield
vanadium. Two species, a chordata and a echinoderm contained vanadium, indicating that
other forms may use vanadium as an oxygen carrier in their vascular systems. The source
of vanadium in sedimentary rock and coals has always been somewhat of a puzzle. It is
possible that such forms as Stickopus mobii may concentrate vanadium and that in depth,
could easily be fixed and held as a constituent of the sedimentary rocks thus formed.
71
Another possibility of the fixation of vanadium under the above conditions is the presence
of hydrogen sulphide which is constantly liberated from muds of mangrove lagoons.
280. . 1918. A possible source of vanadium in sedimentary rocks. American Journal of Science
46: 473-74.
Narrative same as in reference no. 279.
281. Pichard, S. L. and J. R. Paul. 1991. Detection of gene expression in genetically engineered
microorganisms and natural phytoplankton populations in the marine environment by
messenger RNA analysis. Applied Environmental Microbiology 57, no. 6: 1721-27.
A simple method that combines guanidinium isothiocyanate RNA extraction and probing
with antisense and sense RNA probes is described for analysis of microbial gene
expression in planktonic population. Probing of RNA sample extracts with sense-strand
RNA probes was used as a control for nonspecific hybridization or contamination of
mRNA with target DNA. This method enabled detection of expression of a plasmid-
encoded neomycin phosphotransferase gene (nptII) in as few as 10 super(4) Vibrio cells
per ml in 100 ml of seawater. We have used this method to detect expression of the
ribulose-1, 5-bisphosphate carboxylas large-subunit gene (rbcI) in Synechococcus cultures
and natural phytoplankton populations in the Dry Tortugas, Florida. During a 36-h diel
study, rbcL expression of the indigenous phytoplankton was greatest in the day, least at
night(1100, 0300, and 0100 h, and variable at dawn or dusk (0700 and 1900 h). These
results are the first report of gene expression in natural populations by mRNA isolation and
probing.
282. Pitts, R. F. 1936. Clearance values of sucrose and creatinin in the kidneys of the red grouper,
Epinephelus striatus. Carnegie Institution of Washington, Year Book 35: 90-91.
Studies were made on the excretion of urinary creatine nitrogen from the red grouper,
Epinephelus striatus.
283. Plan Development Team, Reef Fish Management Plan South Atlantic Fishery Management Council.
1990. The potential of marine fishery reserves for reef fish management in the U.S.
southern Atlantic, Coastal Resources Division.
Marine fishery reserves (MFRs), areas with no consumptive usage, are recommended as a
viable option for management of reef fisheries in the U.S. southern Atlantic region. MFRs
are designed to protect reef fish stocks and habitat from all consumptive exploitation within
specified geographical areas for the primary purpose of ensuring the persistence of reef fish
stocks and fisheries. Fishery reserves are intended to protect older and larger fishes. This
will benefit reef fisheries by protecting critical spawning stock biomass, intra-specific
genetic diversity, population age structure, recruitment supply, and ecosystem balance
while maintaining reef fish fisheries. The MFR concept is easily understandable by the
general public and possibly more easily accepted than some other management strategies.
Fishery reserves provide some insurance against management and recruitment failures,
simplify enforcement, and have equitable impact among fishery users. Data collection
needs solely for management are reduced and management occurs without complete
information and understanding about every species and interaction. Use of fishery reserves
will establish U.S. leadership in producing model strategies for cooperative international
reef resource management in the Caribbean. Large resident fishes that wander out of
reserved can help maintain certain trophy fisheries. MFR sites with natural species
equilibrium will allow measurement of age, growth, and natural mortality for fisheries
purposes and will provide a basis for other educational, economic, and scientific benefits.
Because there is no fishing within MFRs, impacts of hook and release mortality are
eliminated and the temptation for incidental poaching is reduced. A mixed management
strategy is recommended where 20% of the shelf is MFR, while the remaining 80% is
72
managed for optimal yield by any of several traditional options. Coordinated fishery
reserve efforts in state waters would enhance the benefits of MFRs. Obstacles to fishery
reserves include automatic resistance to new approaches in U.S. marine fisheries,
opposition by some local special interests near proposed reserves, and uncertainty
concerning the size, location, and number of reserves necessary to ensure persistence of the
reef fish fisheries. The incentive for deliberate poaching may be increased within the
reserves; thus, at-sea surveillance and enforcement may be necessary. New artificial reefs
may be needed to replace those lost by inclusion within fishery reserves. Other fishery
management plans should be coordinated to control trolling and other fishing activities
within reserves that may impact reef fishes. The short-term impacts on total harvest caused
by placing fishing habitat into fishing reserves should be compensated for by long-term
fishery benefits. The Dry Tortugas is listed as a potential marine fishery reserve site.
284. Plantier, T. L. 1988. "A comparison of reproductive success in early and late breeding sooty terns
Sterna fuscata in the Dry Tortugas." MS.Thesis, Florida Atlantic University, Boca Raton.
Evidence indicates that earlier-nesting birds are often older, choose preferred nest sites,
and have greater reproductive success than those nesting later. The sooty terns at Bush
Key appear to follow a similar pattern. The first birds arrive at the west end of the
breeding grounds three weeks earlier than birds at the east end and behaviorally appear to
be older and more experienced. The west birds settled in the more desirable habitats (the
west end was cooler than the east end) and laid larger eggs, hatched larger chicks, enjoyed
greater hatchability, fed their chicks at a lower frequency when they were young, and had
greater reproductive success than birds in the east. This was accomplished through a
combination of choosing physically and thermally more favorable habitat, which was more
centrally located, being more persistent incubators and brooders, and, by nesting earlier,
having larger, less-easily eaten chicks by the time avian predators arrived on the island.
285. Plough, H. H. and N. Jones. 1940. Ecteinascidia tortugensis, species Nova; with a review of the
perophoridae (Ascidiacea) of the Tortugas. Papers Tortugas Laboratory 32: 47-60 (issued
Oct. 1939).
Carnegie Institution of Washington Publication Number 517.
During the season of 1936 at the Tortugas Laboratory the senior investigator undertook a
study of the regeneration of pieces cut from the growing stolons of several species of the
family Perophoridae (Ascidiacea Phlebobranchia), which grow in profusion at many places
in the Tortugas area. A new member of the family Perophoridae is described and named
Ecteinascidia tortugensis from its type locality, the Dry Tortugas Key, Florida. It is
shorter than other Ecteinascidia, lies on the ventral side attached along the test, has the
siphons on the dorsal side widely separated and opening in opposite directions, and
possesses a marked secondary loop in the intestine. This species reaches sexual maturity
early in July at the Tortugas, about two weeks later than E. conklini. A brief account of the
development is given. The structure and growth habits of E. tortugensis indicate that it is
intermediate between E. turbinata and Perophora. They suggest a relationship of the
Perophoridae with the Ascidiidae.
286. Porter, J. W. 1977. Pseudorca strandings. Oceans 10, no. 4: 8-16.
This article provides information and observations surrounding the stranding of thirty false
killer whales (Pseudorca crassidens) on the Dry Tortugas Islands near Florida in July
1976. The herd appeared to be protecting an injured male, as evidenced by aspects of
social behavior and agnostic behavior directed at sharks and the author. Among other
suggestions, the author postulates that the injured male was unable to feed due to parasitic
infestation of the ears and consequent impairment of echolocation, which caused the whale
to beach in order to avoid drowning in its weakened state. Other strandings are discussed
in light of the information obtained.
13
287. Porter, J. W., J. F. Battey and G. J. Smith. 1982. Perturbation and change in coral reef
communities. Proceedings of the National Academy of Science 79, no. 5: 1678-81.
Ninety-six percent of surveyed shallow-water Dry Tortugas reef corals died during the
severe winter of 1976-77. Data from skeletal stains indicate that death occurred during the
mid-January intrusion of 14 degree C water onto the reef. In deeper water, community
parameters such as percent cover, species number, and relative abundance showed no
significant change. However, an analysis of competitive interactions at the growing edges
of adjacent colonies reveals a 70% reduction in space competition during this
environmental disturbance. These results can explain high variability in the growth rate of
Floridian reefs and demonstrate the importance of obtaining long-term spatial information
to interpret successional dynamics of complex communities.
288. Porter, J.W., O. W. Meier, L. Chiang and T. Richardson. 1993. Quantification of coral reef change
(Part 2): the establishment and computer analysis of permanent photostations in the Florida
SEAKEYS survey (abs). Proceedings of the Seventh International Coral Reef Symposium,
v.1. Mangilao, Guam: University of Guam Marine Laboratory.
Photostations in five of six locations in the Florida Keys reveal a decline of monitored
coral reef resources during the 1980's when up to 40% of the coral died in some protected
areas. Reductions in the number of species and extraordinary shifts in the pattern of
species abundance occurred in addition to loss of live coral cover. While normally
associated with catastrophic physical disturbances, this coral mortality occurred during a
period without major hurricanes in Florida. Relocatable photostations reveal a multiplicity
of causes for this decline. These include: (1) mortality due to "white band" and "black
band" disease, (2) direct and delayed mortality from "coral bleaching," caused by
abnormally elevated sea temperatures, (3) some mechanical damage, and (4) an increase in
cover by algae. The establishment and sequential analysis of remote sensing data acquired
from permanent photo-stations will be described in detail, as well as limits to the
interpretability of these photogrammetric data.
289. Potthoff, T. and W. J. Richards. 1970. Juvenile bluefin tuna, Thunnus thynnus (Linnaeus), and other
scombrids taken by terns in the Dry Tortugas, Florida. Bulletin of Marine Science 20, no.
2: 389-413.
The identification and seasonal distribution of juvenile scombrids in the waters near the
Dry Tortugas, Florida, are described. Specimens were collected (1960 through 1967) from
regurgitated food of terns. Fishes identified were Thunnus thynnus, Thunnus atlanticus,
Euthynnus alletteratus, Auxis spp., and Katsuwonus pelamis; sizes ranged from 24-146
mm. standard length. For the first time, juvenile bluefin tunas are reported in the Dry
Tortugas region; their presence may indicate that spawning of the species takes place in
the area. Identification methods are discussed, with special emphasis on features of the
axial skeleton and the number of gillrakers over the ceratobranchial bone of the first gill
arch. A method is presented for estimating the standard length of damaged specimens on
the basis of the length of the vertebral column.
290. Powers, P. B. A. 1933. Ciliates infesting the tortugas echinoids. Carnegie Institution of Washington,
Year Book 32: 278-79.
Based on studies conducted on the ciliates of the Tortugas sea-urchins, Echinoida, it was
found that when sea-urchins were infested, they made excellent reservoirs for certain
species of ciliates in which to conduct detailed studies of their internal morphology,
cytoplasmic inclusions and neuromotor apparatus.
291. ——. 1936. Studies on the ciliates of sea-urchins: a general survey of the infestations occurring in
Tortugas Echinoids. Papers Tortugas Laboratory 29: 293-326 (issued Dec. 1935).
Carnegie Institution of Washington Publication Number 452.
74
292. Pratt, H.
293.
294.
With the accumulation of data concerning the ciliate infestations of the alimentary tract of
echinoids, it became of increasing interest to have a complete record of the ciliates
infesting sea-urchins about Tortugas. The writer spent the summer of 1933 at the Tortugas
Laboratory. During this time twelve well-defined species of ciliates were found distributed
among seven species of sea-urchins. The present paper gives a complete account of the
general morphology of these ciliates, as well as a description of the various associations
encountered. Twelve species of ciliates are described which infest the alimentary tract of
seven species of sea-urchins from the region about the Dry Tortugas. Eight of the twelve
are new. Only five of these twelve species show any marked host specificity. The
remaining seven species all show a definite host preference. All of these ciliates are
associated with their host when the latter are found occurring near the tide line. In sea-
urchins taken below a depth of ten fathoms, only four species of ciliates are found :
Cryptochilidium bermudense, Anophrys elongat, Cohnilembus coeci and form M. The
nature of this infestation is one of endocommensalism, there being no present evidence to
indicate pathogenic tendencies for any of its members. Data concerning the geographical
distribution of the ciliates infesting sea-urchins from the localities of Beaufort, North
Carolina, Bermuda and Tortugas have been summarized. It is suggested that the center of
this infestation will be found in the sea-urchins from the region of the Lesser Antilles and
that this infestation has been carried northward along with its host, through the agency of
the Gulf Stream.
S. 1910. Monocotyle floridana, a new monogenetic trematode. Papers Tortugas
Laboratory 4: 1-9.
Carnegie Institution of Washington Publication Number 133.
The genus Monocotyle was established by Taschenber, in 1878, for a worm which he had
found on the gills of the eagle-ray (Myliobatis aquila) at Naples, and which he named
Monocotyle myliobatis. The only other known species is Monocotyle ijimae, which was
discovered in Japan in the mouth of Trygon pastinacea, and described by Goto in 1894.
The worm herein described makes the third member of the genus and was taken from the
gills of the whip-ray (Myliobatis freminvillei) in the Gulf of Mexico and studied at the
Marine Biological Laboratory at Dry Tortugas, Florida. It differs in certain features from
the two other species of the genus, but in the general shape and size of the body, the form
and structure of the suckers, down to the smallest details, and the general arrangement of
the genital organs it shows a close relationship to them, especially to M. ijimae.
. 1910. Parallel transport in tropical trematodes. Science 31: 471-72.
The digenetic trematodes, as well as other internal parasites, have probably in their phyletic
history followed somewhat different rules of descent from those of other animals. The fact
that they live inside of other animals and have a very complex life history must affect their
phyletic development, in that migrations are very much limited, and their structure is very
uniform in the parasites themselves. It is probable that where there are apparently related
species of digenetic trematodes living in widely separated localities, they possess the same
or similar structural features. This does not necessarily indicate that there is a close genetic
relationship between them. These facts are well illustrated by the several species of
digenetic trematodes belonging to the genus Helicometra, which were found in certain
fishes of the Tortugas, Florida, and also occur in the Meditteranean Sea. The species of this
peculiar genus are thus taken as an indication, not that they necessarily bear a close genetic
relationship to one another, but that similar or identical environmental conditions exist for
them in these places, so that they have come to possess in the course of time a structure so
similar that they are included in one and the same genus.
. 1916. The trematode genus Stephanochasmus Looss in the Gulf of Mexico. Parasitology
8, no. 3: 229-38.
75
Two species of the genus Stephanochasmus were found in fishes examined for parasites at
the Dry Tortugas: S. casus Linton and S. sentus Linton. The anatomy of these worms have
| several interesting and unique features. S. casus Linton is described in this article.
295. ————. 1913. The trematode parasites of the loggerhead turtle. Science 37: 264-65.
The studies of trematode parasites of the loggerhead turtle in the Mediterranean and the
Gulf of Mexico, Dry Tortugas are briefly discussed and compared. Nineteen species
occurred in the turtles of the Gulf of Mexico. Eight of these species also occur in the
Mediterranean Sea. The most numerous trematode occurring in the Dry Tortugas is
Cymatocarpus undulatus. A more detailed discussion is found in the Archives de
Parasitologie article by Pratt.
296. ———. 1912. Trematodes of the Gulf of Mexico. Verhandlungen des VIII Internationalen
Zoologen- Kongresses, 780-781. Jena, G. Fischer.
This is a discussion of the article written by Pratt in no. 133 Carnegie Institution of
Washington, listing the trematodes collected at the Dry Tortugas and the hosts they live in.
297. ————. 1916. Trematodes of the loggerhead turtle (Caretta caretta) of the Gulf of Mexico. Archives
De Parasitologie: 411-27.
Five species of trematodes are discussed in this paper. Reference is made to the studies of
Linton (1910), as well as studies made in the Mediterranean Sea. The five trematode
species were found in turtles captured on Loggerhead Key in the Dry Tortugas: Wilderia
elliptica, Pachypsolus tertius, and Rhyditodes secundus, Pelsiochorus cymbiformis and
Cymatocarpus undulatus.
298. Raim, A. W., W. Cochran and R. D. Applegate. 1989. Activities of a migrant merlin during an
island stopover. Journal of Raptor Research 23, no. 2: 49-52.
Activities of a radio-tagged merlin (Falco columbarius) which was trapped and identified
as an adult female by George Allex and Daniel D. Berger, were observed from 10-16 April
1977 on Loggerhead Key.
299. Reighard, J. 1908. An experimental field-study of warning coloration in coral-reef fishes. Papers
Tortugas Laboratory 2: 257-325.
Carnegie Institution of Washington Publication Number 103.
This paper embodies a search for the biological significance of the conspicuousness which
it attempts to show characterizes many of the coral-reef fish of the Tortugas region. After
showing that this conspicuousness is not a secondary sexual character and that it serves
neither for protective nor aggressive resemblance, its value as a warning character is
subjected to experimental test. Experimental evidence is presented to show that the gray
snapper, the commonest predaceous fish, discriminates certain colors, forms associations
with rapidity, and retains these for a considerable time (memory). If any of the coral-reef
fishes possess a combination of consicuousness with such unpleasant attributes as render
then unpalatable, the gray snapper should have learned to avoid them at sight and their
conspicuousness would then have a warning significance. It is shown that when atherina,
an inconspicuous fish which serves normally as the food of the gray snapper, is given an
artificial warning color and at the same time rendered unpalatable, it is after a brief
experience, no longer taken as food by the gray snapper. Artificially colored atherinas thus
come to have a warning significance for the gray snapper and are avoided, even when not
unpalatable, although normal atherinas are still readily eaten. The conclusion is thus
reached that the existence of a warning coloration or of warning conspicuousness in coral
fishes is easily possible. The conclusion is reached that the conspicuousness of coral-reef
fishes, since it is not a secondary sexual character and has no necessary meaning for
protection, aggression, or as warning, is without biological significance. The coral-reef
76
fishes have no need of aggressive inconspicuousness because their food consists of
invertebrates, chiefly fixed. They have no need of protective inconspicuousness because
the reefs and their agility afford them abundant protection. Selection has therefore not
acted on their colors or other conspicuous characters, but these have developed in the
absence of selection and through internal forces. They are the result of race tendency
unchecked by selection .
300. . 1907. The photography of aquatic animals in their natural environment. Bulletin of the
United States Bureau of Fisheries 27: 41-68.
This paper describes the photography of aquatic organisms in their native environment and
under normal conditions by carrying the camera into the field. Photos and diagrams are
provided for cameras and apparatuses that remain above the surface of the water and
cameras that are submerged.
301. Reynolds, J. E. III and J. C. Ferguson. 1984. Implications of the presence of manatees, Trichechus
manatus near the Dry Tortugas Islands, Florida USA. Florida Scientist 47, no. 3: 187-89. |
Two West Indian manatees (Trichechus manatus) were observed 61 km northeast of the
Dry Tortugas Islands, a location not normally considered to be part of the species range. )
When spotted, the animals were swimming in a soutwesterly direction, away from Florida.
Observations such as this, of manatees far from freshwater, raise the question of whether
manatees require regular access to freshwater for osmoregulation, as suggested in the
literature.
302. Reynolds, J. E. III and J. C. Steinmetz. 1983. Dry Tortugas: products of time. Sea Frontiers 29, no.
2: 66-75.
This article discusses the general formation of the islands and their history.
303. Richards, O. W. 1934-1936. Growth studies in the ascidian, Phallusia nigra, and hermit crab,
Caenobita clypeatus. Carnegie Institution of Washington, Year Book
Note: published as follows 1934, v. 33, p. 261; 1936, v. 35, p. 92.
The early growth and development of the ascidian Phallusia nigra was recorded over time
using motion and still photography. Claw size ratios to body size of the hermit crab,
Coenobita clypeatus, were examined by correlation methods.
304. Ricklefs, R. E. and S. C. White. 1981. Growth and energetics of chicks of sooty tern, Sterna fuscata
and common tern, Sterna hirundo. Auk 98, no. 2: 361-78.
The energy budgets of chicks of the common tern (Sterna hirundo) were measured on
Great Gull Island, New York. Also measured were the sooty tern (S. fuscata) on the Dry
Tortugas, Florida The respiratory energy requirement was determined by measuring
oxygen consumption in a closed system. The growth energy requirement was calculated
from the lipid and protein contents of a series of chicks spanning the range between
hatching and fledging. Energy budgets calculated for the two species differed in several
ways. (1) Maintenance metabolism was lower in the sooty tern owing to its warm
environment. (2) Sooty terns allocated more of their energy intake to lipid accumulation
from an earlier age. (3) In the sooty tern, the allocation of energy to growth initially was
high, but its absolute amount decreased steadily throughout the growth period. In the
common tern, both growth and maintenance energy allocations increased rapidly during the
first half of the development period. (4) In sooty tern chicks energy metabolism
approached its maximum rate (135 kJ/day) by the end of the first third of the development
period, after which it leveled off. In the common tern, energy metabolism increased from
about one-quarter of its maximum during the first five days after hatching to its maximum
of 200kJ/day during the third week of the postnatal development period. Although these
observations support the hypothesis that slow growth in pelagic seabirds is selected to
Vi
reduce the energy requirement of the chick, the energy budgets also suggest that a doubling
of the growth rate by the sooty tern would increase the maximum energy requirement of the
chick by only 20% and the total feeding requirement of the adult by only 5%. Moreover,
the levels of water in muscles suggest that the sooty tern develops mature function earlier
than does the common tern, which in itself might be sufficient to account for the slower
growth of the first species.
305. Ricklefs, R. E. and S. C. White-Schuler. 1978. Growth rate of the brown noddy on the Dry Tortugas
Florida USA. Bird-Banding 49, no. 4: 301-12.
Growth rates within seabird species can vary with locality, season, and year. In this study
noddy tern checks captured on Bush Key, Dry Tortugas, June, 1972, were weighed and
measured. Growth increments were used to calculate a composite wing length growth
curve to estimate the ages of chicks. A logistic curve was fitted to describe the relationship
between weight and age. Growth constants of the fitted curve (growth rate K = 0.153,
asymptote A=160 grams, and age at inflection ti=14.0 days) were similar to values reported
for the brown noddy on Kure Island and Manana Island, Hawaii. Also reported are outer
primary and rectix lengths and body temperatures of nestlings and adults.
306. Riley, G. A. 1938. Study of the plankton in tropical waters. Carnegie Institution of Washington
Year Book 37: 98.
The small quantity of plankton in tropical waters as contrasted to higher latitudes is
investigated, and when compared to a similar survey underway in Long Island, N.Y., the
indication is that chlorophyll and plant pigments are one-twenty-fifth the amount found in
New York.
307. Riska, D. E. 1986. An analysis of vocal communication in the adult brown noddy, Anous stolidus.
Auk 103, no. 2: 359-69.
The author analyzed vocal signals of marked adult Brown Noddies (Anous stolidus)
throughout their nesting season in the Dry Tortugas, Florida from 1979 to 1982. The basic
unit of the adult repertoire is a wide-band click, less than 4 msec duration, ranging in
frequency from 200 to 3,300 Hx. He identified nine temporal arrangements of these clicks,
which form the notes of the calls. These calls differ little in frequency range, but they
differ in the mean frequency of the most intense sound energy band, in note duration, in the
number of clicks per note, and in internote interval. These calls are used in different
contexts, which sometimes overlap. Frequency, note duration, and length varied among
individuals for some calls. No tonal elements characteristic of calls of brown noddy
nestlings remain in the adult repertoire.
308. ———. 1986. "Communication behavior of the brown noddy (Anous stolidus) and sooty tern
(Sterna fuscata), Dry Tortugas, Florida (vocalizations, laridae, signals, colonial,
breeding)." Ph.D. Dissertations, University of California at Los Angeles.
The basic unit of the adult repertoire is described as a wide-band click, less than 4 msec
duration, in the frequency range 200 to 3300 Hz. Nine calls differ in temporal
arrangements of clicks, mean frequency of the most intense sound energy band, note
duration, number of clicks per note, and inter-note interval. Frequency, not duration, and
inter-note interval do not differ between sexes. The nestlings of the brown noddy produce
three structurally different vocalizations within one day after hatching. Postures of chicks
and contexts in which these signals are used differ. The repertoire is composed of
frequency-modulated tonal elements and broad-band bursts of sound with little
resemblance to the adult repertoire. Juvenile bush-nesting noddies begin flying when 40-
48 days old, after which they are still fed at their nests. Adult noddies accept a substituted
nestling differing from their own in size, color and plumage stage, up to at least 20 days
post-hatching. The adult Sooty Terns produce eight structurally different vocalizations,
78
and nestlings produce three, in the frequency range 300-7000 Hz. Postures differ for each
call, but contexts in which these are used overlap. The range of frequencies in which
young birds call extends higher than that of adults, but the frequency-modulated tonal
elements characteristic of nestling vocalizations remain complex in the adults.
309. Rivas, L. R. 1951. Preliminary review of the western North Atlantic fishes of the family
310. Roberts,
311.
Roberts,
Scombridae. Bulletin of Marine Science of the Gulf and Caribbean 1, no. 3: 213-30.
This paper brings up to date the taxonomy of the western North Atlantic mackerels and
tunas. In addition to a key to the genera and species, a complete synonymy, a diagnosis
and pertinent comments are given under each species.
H. H., L. J. Rouse Jr., N. D. Walker and J. H. Hudson. 1982. Cold water stress in Florida
Bay and northern Bahamas, a product of winter cold air outbreaks. Journal of Sedimentary
Petrology 52, no. 1: 145-55.
During January 1977 three consecutive cold fronts crossed south Florida and the northern
Bahamas which depressed shallow-water temperatures below the lethal limit for most reef
corals. Digital thermal infrared data acquired by the NOAA-5 meteorological satellite, in
situ water temperatures, and meteorological data were used to study the thermal evolution
of Florida Bay and Bahama Bank waters. The third and most important frontal system
depressed Florida Bay water below 16 degrees C, a thermal stress threshold for most reef
corals, for 8 days. Coral mortality at Dry Tortugas was up to 91 percent during the 1977
event. Coral and fish kills were also reported from other parts of the Florida Reef Tract
and northern Bahamas. Study results show that cold-water stress conditions can exist over
vast shallow-water areas and have residence times of several days.
H., H. Lawrence , J. Rouse Jr. and N. D. Walker. 1983. Evolution of cold water stress
conditions in high-latitude reef systems: Florida Reef Tract and the Bahama Banks.
Caribbean Journal of Science 19, no. (1-2): 55-60.
Thermal depression of shallow bank and bay waters accompanying the passage of severe
cold fronts can stress high latitude coral reef systems, such as those of the Florida Reef
Tract and northern Bahama Banks. Laboratory and field experiments suggest that
sustained temperatures below 16 degrees C are detrimental to most reef-building corals.
Time-series satellite imagery provides a data base for assessing the thermal variability of
waters interfacing with reef systems. Digital thermal infrared data acquired by the NOAA-
5 meteorological satellite were used to study thermal evolution of Florida Bay and Bahama
Bank waters during a succession of three cold-air outbreaks (January 1977). These studies
indicate that the temperature of subtropical bank and bay waters is subject to depression
below 16 degrees C accompanying the outbreak of unusually cold air. This superchilled
water can have a residence time of days. The cooling process creates water masses that are
out of density equilibrium with warmer ocean water. Offshelf movement of the cold, dense
water occurs at particular sites, as shown by time-series satellite data. The absence of coral
reefs opposite tidal passes in the Florida Keys is attributed to this process, which has
probably limited development of the entire reef tract.
312. Robertson, W. B. Jr. 1978. Species of special concern sooty tern. Birds 2, no. Edited by H. Kale:
89-90.
A description, range, and habitat of the sooty tern are given along with its life history and
ecology at the Dry Tortugas. Its classification is based not on its abundance, but it is
because the Dry Tortugas colony is a major Florida wildlife resource. Aside from the
Tortugas no other location in Florida is suitable for sooty tern nesting. This colony affords
a means of monitoring the general health of offshore Gulf waters of southern Florida.
gS
S113), . 1964. The terns of the Dry Tortugas. Bulletin of the State Museum , Biological Science 8,
no. 1: 1-95.
New information from unpublished sources and from published records hitherto
overlooked permit a re-evaluation of the history of the Dry Tortugas and of the terns that
inhabit them. The geography and ecology of the 11 keys that have variously comprised the
group since it was first mapped in the 1770's are described and their major changes traced.
The recorded occurrences of the seven species of terns reported nesting on the Keys are
analyzed in detail. The sooty tern colony has fluctuated from a low of about 5,000 adults
in 1903 to a reported peak of 190,000 in 1950; for the past four years it has remained
steady at about 100,000. The brown noddy population, which reached a peak of 35,000 in
1919, was reduced by rats to about 400 adults in 1938; it is in the neighborhood of 2,000
today. A colony of 150 to 450 roseate terns has nested in most years from 1917 to the
present. About 500 least terns nested regularly trom 1918 to 1932, then unaccountably
dwindled to a few pairs by 1937 and shortly afterward disappeared. Royal and sandwich
terns nested abundantly in the mid-19th century, and a colony of royals may have existed as
late as 1890. Both species are believed to have been extirpated from the Tortugas by
egging. No verifiable evidence exists for the nesting of the common tern, which has been
reported several times. The black noddy, first reported for the continental United States at
Dry Tortugas in 1960, has been found there each summer since.
314. . 1969. Transatlantic migration of juvenile sooty terns. Nature 222: 632-34.
From 1959 to 1968, 70,000 adult and 130,000 juvenile sooty terns (Sterna fuscata) were
banded at Bush Key, Dry Tortugas, Florida. By December 1968, 29 juveniles were
recovered in West Africa. It appears that the primary biological function of the
transatlantic migration is to avoid intraspecific competition and this adaptive value
becomes evident when the migration of juveniles is seen in the context of the rigidly
structured sooty tern population. It may be evidence for a successful evolutionary
mechanism.
315. Robertson, W. B. Jr. and B. Given. 1980. Ruddy quail dove Geotrygon montana again at Dry
Tortugas Florida USA. Florida Field Naturalist 8, no. 1: 23-24.
About noon on December 15, 1977, a cold day with severe northwesterly squalls, Given
found and photographed a large, reddish dove on the second tier of Fort Jefferson, Dry
Tortugas, Florida. This record is the fifth report of the species from Florida and the second
from Dry Tortugas.
316. Robertson, W. B. Jr. and C. R. Mason. 1965. Additional bird records from the Dry Tortugas.
Florida Naturalist 38: 131-38.
Sprunt (1962-63) summarized what was known about the occurrence of birds at the Dry
Tortugas through the Summer of 1962. In this paper the authors report on recent bird
records up to April 1965. Comments relate to 12 species new to the list or those known
from either one or two records. Sprunt listed 227 species of birds for the Tortugas, the
authors add 12 to bring the total to 239 species.
317. Robertson, W. B. Jr. and L. C. Below. 1975. A red-headed woodpecker at Dry Tortugas. Florida
Field Naturalist 2, no. 1: 20.
On May 5, 1973 Mr. and Mrs. G.H. Perbix of Cincinnati and Mrs. Below, members of the
tern-banding party then at Dry Tortugas, visited Loggerhead Key and at once noticed an
adult red-headed woodpecker (Melanerpes erythrocephalus) in the large Australian Pines
(Casuarina equisetifolia) near the dock. We find only one other report of the red-headed
Woodpecker at Dry Tortugas. Howell (1932:308) wrote that the species was unknown in
the Florida Keys "...except for a single occurrence on the Tortugas - a bird seen there on a
number of days early in June." The red-headed woodpecker is not known to occur outside
80
the United States but the present record inevitably raises the question: was the bird
migrating across the Gulf or was it merely a vagrant?
318. Robertson, W. B. Jr. D. R. Paulson and C. R. Mason. 1961. A tern new to the United States. Auk
78: 423-25.
This note provides a description of the black noddy, Anous tenuirostris collected at Dry
Tortugas. This is the first of this species collected in the United States. Two specimens
were taken from Bush Key during July 1960. The bird occurs nearly world-wide in the
warmer seas, but is absent from most of the Atlantic Ocean north of the equator and most
of the Caribbean Sea.
319. Robinson, A. H. 1976. Marine, island and coastal parks in the United States National Park system:
A review and progress report in 1975. International Conference on Marine Parks and
Reserves., pp. 226-27. Gland, Switzerland: IUCN.
This paper provides a basic introduction to critical marine habitats and the planning and
management of marine parks and reserves, including interpretation and environmental
education in marine parks. Progress in the creation of marine parks and reserves is
reviewed, and a special report on marine park systems in the Pacific region is included.
Fort Jefferson National Monument is discussed as the first underwater preserve established
in the United States.
320. Schaeffer, A A. 1925. Experiments on the influence of temperature and dilute and concentrated
sea-water on ameboid movement. Bulletin of the Ecological Society of America: 11.
The reactions of various species of amoebas to different concentrations of sea-water have
been used during the past several years at Tortugas as important aids in the identification
and fixation of species. The rate of movement of several species was studied in various
concentrations of sea-water indicating that the optimal concentration of sea-water is below
the norm in every case when measured by the rate of cell-coordinated movement.
321. ————. 1926. Taxonomy of the amebas: with descriptions of thirty-nine new marine and
freshwater species. Papers Tortugas Laboratory 24: 1-116.
Carnegie Institution of Washington Publication Number 345.
The purpose of this report is to set forth a description of 39 new species and 11 new genera
of amebas (Amoebaea), and to propose a preliminary system of classification of the
amebas, based on their general morphology. General observations on structure,
physiology, distribution, and methods of investigation are provided. The changes of form
which amebas undergo is a fundamental morphological characteristic of amebas, and forms
the basis of a natural classification. For the purpose of quickly recognizing a species other
characteristics are more valuable, such as the nucleus, vacuoles, crystals, resistance to
dilutions and concentrations of sea-water, etc. A brief discussion of these characteristics
with reference to specific descriptions is given along with colored drawings and
photographic text-figures.
322. Schmitt, W. L. 1924-1932. Systematic-ecologic studies of the decapod crustacea. Carnegie
Institution of Washington, Yearbook.
Note: published as follows: 1924, v.23, p. 200-201; 1925, v.24, p. 230-231; 1930, v.29, p.
34371931, v:30) pe 3892 193825v. 37, p. 279:
Very striking color characteristics/variations affecting chela, and often the appendages, are
noted among snapping shrimp, Synalpheus, and giant isopod crustaceans. Bathymetric
distribution of decapods are investigated.
323. Schnell, G. D. 1974. Flight speeds and wing beat frequencies of the magnificent frigate bird. Auk
91, no. 3: 564-70.
81
Wingbeat frequencies and flight speeds of magnificent frigatebirds were recorded with a
Doppler radar in the Dry Tortugas, Florida. The flapping rate averaged 2.84 beats per
second (SD 0.14) and was not significantly correlated with flight speed, providing further
evidence that the birds’ wingbeat frequency is essentially constant within species. The
flapping rate is somewhat higher than predicted from the theory of mechanical oscillators
when the distance from the end of the wing to the first articulated joint is used as an
estimate for the average effective wing length. Flight speeds of birds in a flat calm
averaged 22.55 mph. The highest average ground speed of 30.17 mph was obtained from
frigatebirds flying in a 6 to 8 mph wind, and the lowest of 16.00 mph for birds flying into
the 65 mph wind. Airspeeds were greater for frigatebirds flying into the wind than for
those moving across or with the wind.
324. Schreiber, R. W., W.B. Robertson Jr. and T. Bellow. 1976. Nesting of brown pelicans, Pelecanus
occidentalis, on the Dry Tortugas, Florida. Florida Field Naturalist 3, no. 2: 47-48.
On June 14, 1974 Bush Key, Dry Tortugas Ted Bellow and C. Winegarner found 5 brown
pelican nests about 12 feet above ground in the white mangroves (Laguncularia racemosa)
along the north shore. Nineteenth-century records of pelicans breeding on the Dry
Tortugas are ambiguous. ...on the Tortugas (1860) it thus appears that a few pairs did breed
on the Tortugas in the mid-1900's, but by late in the century none did so. Our record is the
first reported nesting of this species in the 20th century on the ornithologically well-known
islands (Robertson and Mason, 1965). Three of the nests found in 1974 contained two
eggs each, one nest was empty, and the fifth was not checked.
325. Schroeder, P. B. and J. H. Davis. 1971. Ecology vegetation and topography of the Dry Tortugas
updated to 1970. Quarterly Journal of the Florida Academy of Science (Supp! 1): 12-13.
The half-dozen islets of the Dry Tortugas have been ecologically studied periodically since
the turn of the century. In November and February a year ago, a field party from the
University of Miami made a topographical and vegeational study of several of these keys.
The pertinent information gathered at that time is now available and provides continuity
with the studies of Millapaugh (1907), Bowman (1918) and Davis (1942). The keys
studied have changed from barren coral and sand to substantial islets largely covered with
vegetation. The configuration of one of these has been completely altered. All the keys
have been changed considerably in shape. Vegetational communities have shown similar
changes and maturity. Mangrove areas (red and white) have become established and
enlarged. Australian pines and other exotics, introduced to Loggerhead Key, have spread
over much of the island and now are found on Bush Key.
326. Scott, W. E. D. 1890. On birds observed at the Dry Tortugas, Florida, during parts of March and
April, 1890. The Auk: A Quarterly Journal of Ornithology 7, no. 4: 301-14.
The list of birds observed at the Tortugas includes eighty species, fifty seven of which were
land birds. The author states that no land birds breed on any of the keys group, and that the
stay of any land bird is of very short duration.
327. Seaman, W. Jr. and D. Y. Aska. 1974. Research and information needs of the Florida spiny lobster
fishery. State University System of Florida Sea Grant Program, Miami FL. 64 pgs.
In response to a number of fishermen in South Florida, the State University System of
Florida Sea Grant Program became involved in research on the spiny lobster, Panulirus
argus. When additional research needs were expressed, Florida Sea Grant decided to
become better informed on the subject, and evaluate its potential for service to the persons
dependent on this fishery resource. A meeting of persons and organizations involved in the
biology and/or utilization of the spiny lobster fishery in Florida was called to identify
broadly the problems and information needs of persons dependent on the spiny lobster
82 :
resource, to assess existing sources of information and their possible applications, and to
identify priorities and actions needed to resolve user problems.
328. Shinn, E. A., J. H. Hudson, R. B. Halley and B. Lidz. 1977. Topographic control and accumulation
rate of some Holocene coral reefs: South Florida and Dry Tortugas. Proceedings, Third
International Coral Reef Symposium, RSMAS, Univ. of Miami, Coral Gables FL. p.1-7.
Core drilling and examination of underwater excavations on 6 reef sites in south Florida
and Dry Tortugas revealed that underlying topography is the major factor controlling reef
morphology. Carbon-14 dating of coral recovered from cores enables calculation of
accumulation rates. Accumulation rates were found to range from 0.38 m/1000 years in
thin Holocene reefs to as much as 4.85 m/1000 years in thicker buildups. Cementation and
alteration of corals were found to be more pronounced in areas of low buildup rates than in
areas of rapid accumulation rates. Acropora palmata, generally considered the major reef
builder in Florida, was found to be absent in most reefs drilled. At Dry Tortugas, the more
than 13-meter thick Holocene reef did not contain A. palmata. The principal reef builders
in this outer reef are the same as those which built the Pleistocene Key Largo formation,
long considered to be a fossilized path reef complex.
329. Shinn, E. A. 1984? Geologic history, sediment, and geomorphic variations within the Florida Reef
Tract. Advances in reef sciences, abstracts and schedule of presentations: a joint meeting
of the Atlantic Reef Committee and the International Society for Reef Studies , 113-14.
Miami, Florida: University of Miami.
A combination of core drilling, high resolution seismic profiling, and constituent particle
analysis reveal these major aspects of Holocene reef development and sediment
distribution within the Florida reef tract: (1) reef distribution and shape are controlled by
underlying Pleistocene limestone topography; (2) accumulation of sand and rubble occurs
in forereef and backreef areas; and (3) composition of sediment and coral distribution are
controlled by the reef tract trend relative to prevailing wind and exposure to Gulf of
Mexico water.
330. Shinn, E. A., B. H. Lidz, R. B. Halley, J. H. Hudson and J. L. Kindinger. 1989. Reefs of Florida
and the Dry Tortugas: Miami to Key West, Florida, July 2-7, 1989 . 28th International
Geological Congress. Field Trip Guidebook (American Geophysical Union), no. T176.
Washington, D. C.: American Geophysical Union.
This field guide concentrates on explaining the distribution of Holocene coral reefs, the
relationship between topography and Holocene sea-level rise, and the compositional and
thickness variation of sediments produced in and adjacent to the reefs. A discussion and
speculation of the future of the reefs under a stable sea, and a lowered sea-level is included.
Also attached is a key to the Stony Corals of the Florida Keys and Dry Tortugas, a species
list, illustrations of geologic cross-sections, aerial and underwater photographs of reefs and
coral.
331. Shoemaker, C. R. 1934. Two new genera and six new species of Amphipoda from Tortugas. Papers
Tortugas Laboratory 28: 245-56 (issued Nov. 1933).
Carnegie Institution of Washington Publication Number 435.
New genera and species of Amphipoda are described from specimens collected at the
Tortugas, including Socarnes concavus, Gitanopsis tortugae, Heterophilas seclusus,
Pontogeneia longleyi, Ampithoe divursia, Leucothoides pottsi (new species); and
Heterophlias, Leucothoides (new genera).
332. Silberman, J. D., S. K. Sarver and P. J. Walsh. 1994. Mitochondrial DNA variation and population
structure in the spiny lobster Panulirus argus. Marine Biology 120: 601-8.
Adult spiny lobsters (Panulirus argus) were collected from nine locations including the
83
Tortugas, throughout the tropical and subtropical northwest Atlantic Ocean and examined
for mitochondrial DNA (mtDNA) variation. 187 different mtDNA haplotypes were
observed among the 259 lobsters sampled. Haplotype diversity was calculated to be 0.986
and mean nucleotide sequence-diversity was estimated to be 1.44%; both of these values
are among the highest reported values for a marine species. Analysis of molecular variance
(AMOVA) and phenetic clustering both failed to reveal any evidence of genetic structure
within and among populations of P.argus The present data are consistent with high levels
of gene flow among populations of P.argus resulting from an extended planktonic larval
stage and strong prevailing ocean currents.
333. Smayda, T.J., Y. Shimizu, C. R. Tomas and D. G. Baden. 1993. The influence of phosphorus
source on the growth and cellular toxin content of the benthic dinoflagellate Prorocentrum
lima. Toxic Phytoplankton Blooms In The Sea., 565-70.
The relationship between toxin content and nutritional status of the toxic marine
phytoplankton species Prorocentrum lima was examined in a clonal culture isolated from
the Dry Tortugas, Florida, grown with inorganic phosphate and glycerol phosphate
enriched media. Growth, alkaline phosphatase activity and okadaic acid content were
measured. Phosphate enriched cultures exhibited rapid growth rates(0.75 div/d), moderate
terminal densities of 134,779 cells/ml and low alkaline phosphatase activity (<14
fg/cell/min). Cells grown with glycerol phosphate had lower growth rates, between 0.16
and 0.45 div/d, but higher maximal densities, >200,000 cells/ml, and had alkaline
phosphatase activity an order of magnitude greater than those grown in inorganic
phosphate. When comparing toxin levels at 20 and 30 days, cells grown on the organic
phosphate enrichments had consistently higher per cell values (11.2 and 14.2 pg/cell,
respectively) than those with inorganic phosphate (7.5 and 8.9 pg/cell), respectively).
Phosphorus source effected growth, maximal densities, and okadaic acid content of P.
lima. ;
334. Smith, H. G. 1937. Contribution to the anatomy and physiology of Cassiopea frondosa. Papers
Tortugas Laboratory 31: 17-52 (issued July 1936).
Carnegie Institution of Washington Publication Number 475.
This research was undertaken to extend our previous scanty knowledge on the physiology
of feeding and digestion in the Scyphozoa. Cassiopea was selected as the experimental
material for two reasons, it is a member of the Rhizostomeae in which the mode of feeding
is particularly interesting owing to the sub-division of the mouth, while certain species,
including the one studied, possess zooxanthellae. Other aspects of the structure and
physiology of species of this genus have been extensively studied, notably at the Tortugas
Laboratory, by Mayer and others. It was originally intended to work on C. xamachana,
which was very abundant at one time in the moat at Fort Jefferson. Recent changes in the
conditions in the moat, the result of silting up, have caused the complete disappearance of
the species although from this locality. In the absence of this species, C. frondosa was
investigated and this although less hardy than C. xamachana, proved satisfactory material.
Experiments have been made on the effect of starvation in light and in darkness on the
medusae. In light, specimens were kept alive for 15 days, and in darkness for 7 days.
Numerous algae were ejected by way of the gastric filaments and plaited membranes at the
base of the filaments and the medusae became brown in color. They also shrank
considerably in size. The effect of the zooxanthellae on phosphorus excretion has been
studied, the amount of phosphorus in the sea-water surrounding one specimen being
reduced to zero within 24 hours. Finally, feeding, digestion and symbiosis in C. frondosa
have been discussed. It has been suggested that the variation in pH in the coelenteron
affects the activity of the jellyfish. The association with zooxanthellae is probably similar
in nature to that which occurs in the Madreporaria.
84
335. South Florida Area : Synthesis of available biological, geological, chemical, socioeconomic and
336. Spence,
cultural resource information. 1990. OCS Study, MMS 90-0019. U.S. Department of the
Interior, Minerals Management Service, Atlantic OCS Regional Office.
This study summarizes the available biological, geological, chemical, and socioeconomic
information in south Florida in relation to the potential effects of offshore gas and oil
exploration and development. The synthesis will help Federal and state policy makers
make informed decisions about future lease offerings and environmental restrictions on
offshore oil and gas operations. The Dry Tortugas is included as part of the South Florida
Reef Tract. In summary it would be very difficult to protect the mangroves, reefs, seagrass
beds and their associated assemblages from large oil slicks. Severe weather would make it
impossible. The Dry Tortugas experienced an oil spill from the beaching of Brother
George in 1964. Birds were killed . Some coral may have been killed around the
Tortugas from the 3,100 barrel spill, but it did not affect other areas further to the east of
the site. If a large oil spill did occur here it would take 100+ years for the oldest coral
heads to regrow and achieve the same level of pre-spill structural complexity. The effects
of an oil spill on other flora and fauna of the Florida Reef Tract can only be guessed.
J., and O. W. Richards. 1940. Native cellulose in the ascidian Phallusia nigra. Papers
Tortugas Laboratory 32: 163-67 (issued Sept. 1940).
Carnegie Institution of Washington Publication Number 517.
Many organic compounds of high molecular weight are readily identifiable from the
characteristics of the X-ray diffraction diagrams. Cellulose and its derivatives have been
extensively examined by X-ray diffraction methods in the search for a complete solution of
the structure and crystallite arrangement of the cellulose molecule. From the analytical
standpoint, X-ray diffraction diagrams not only confirm the initial chemical identification
of cellulose by Schmidt (1845), but they also show the presence of crystallites and their
orientation. The Phallusia nigra was collected in the moat of Fort Jefferson and the tunic
was removed on return to the Tortugas Laboratory. The result, namely, the recognition of
native cellulose and the preferred orientation of the crystallites in Phallusia nigra, is
naturally anticipated from previous observations on other ascidian tests. This method
provides a useful analytical "tool" for use in zoological investigation.
337. Sprunt, A. Jr. 1963. Birds of the Dry Tortugas. Florida Naturalist: 22-26, 52-53.
338. ——.
339, ———_.
340. ———.
341. ———.
This is a continuation of the listing from the 1962 series on listings of birds of the Dry
Tortugas.
1962. Birds of the Dry Tortugas 1857-1961. Florida Naturalist 35: 35-40, 82-85, 129-32.
A brief discussion of the history of bird studies of the Dry Tortugas is given. Special
attention is paid to the migratory birds passing through the Dry Tortugas in hope of
shedding light on trans-Gulf migration.
1947. Blizzard of birds: the Tortugas terns. National Geographic Magazine February:
213-30.
This article gives a history of the Tortugas terns up to 1947. Boobies and noddies are
included also.
1950. Bridled tern, Sterna a. Melanoptera, taken at Dry Tortugas. Auk 67, no. 4: 514.
This article provides an account of the first Sterna melanoptera recorded at the Dry
Tortugas, and the fifth specimen recorded in Florida.
1950. A list of birds of the Dry Tortugas Keys, 1857-1949. Florida Naturalist 23: 49-60,
73-78, 105-11.
342.
343.
85
A listing of the birds of the Dry Tortugas is given. Land birds-pigeons through vireos-
including warblers through sparrows and water birds.
. 1951. Some observations on the fall migration at Dry Tortugas. Auk 68: 218-26.
The author arrived at the Tortugas following a hurricane August 26-27, which seemed to
have no effect on the Tortugas, or the birds there. He found the birds to be in good
physical condition, with no signs of exhaustion. Birds were tame and could be approached.
A listing of the birds sighted is given.
. 1948. The tern colonies of the Dry Tortugas keys. Auk 65: 1-19.
The first post-war (1945-46) status report on tern populations inhabiting the Keys of the
Dry Tortugas is presented in this paper. A brief history on population counts dating back
to 1832 by Audubon is given, as well as a description, mostly vegetative, on Keys utilized
by terns for nesting activities. Tern springtime arrival and summer departure are discussed,
along with numbers of eggs produced, nesting locations and tern behavior. Based on the
square-yard unit system, it was determined that the population count for Sooties was
97,200, while a count of 550 was found for the noddies by numbers of nests. The tern
populations have suffered virtually no damage during the occupation of the islands by
naval forces. Aside from weather, predation by natural enemies includes sand-crabs and
man-o'-war-birds. The tern colonies appeared safe, but certain topographical changes, such
as the recent increase in vegetation may be problematical.
344. Stevenson, J. O. 1938. The tern colonies of Dry Tortugas. Bird-Lore 40, no. 5: 305-9.
This article describes briefly the history of the tern colonies of the Dry Tortugas. The
author visited the Tortugas on May 24, 1937, two years after a hurricane swept through the
islands destroying Bird Key, the historic breeding grounds for thousands of sooty and
noddy terns.
345. Steward, F. C. 1940. The growth of Valonia ventricosa J. Agardh and Valonia ocellata Howe in
culture, with a note on the sap composition of Valonia ocellata Howe at Tortugas. Papers
Tortugas Laboratory 32: 85-98 (issued Oct. 1939).
Carnegie Institution of Washington Publication Number 517.
So much physiological work has been done using species of Valonia that their mode of
development has special interest. Living material of V. ventricosa and V. ocellata was
collected at the Dry Tortugas, Florida. These species were chosen because of the
difference in their morphology. Valonia ventricosa J. Agardh and V. ocellata Howe have
been kept alive for over two years from their original collection. Vesicles of considerable
size (V. ventricosa) and with all the characteristics of the plant in nature have been grown
attached to a suitable substratum. The development of the vesicle and rhizoids from
aplanospores is illustrated by a series of photographs. V. ventricosa also produces
filaments which penetrate the substratum and from which close clusters of vesicles arise as
they do in the normal habitat. The appearance of the aplanospore and growing vesicle
between crossed Nicols is described and its bearing on the structure of the wall indicated.
Valonia ocellata produces pear-shaped vesicles, cylindrical rhizoidal processes (which it is
shown may become long and branched), and apparently proliferated masses composed of
small cellular segments. The growth and development of all these structures from
aplanospores, or the product of "segregative division" have been observed and are
recorded by photographs.
346. Steward, F. C. and J. C. Martin. 1937. The distribution and physiology of Valonia at the Dry
Tortugas, with special reference to the problem of salt accumulation in plants. Papers
Tortugas Laboratory 31: 87-110 (issued Oct. 1936).
Carnegie Institution of Washington Publication Number 475.
86
This paper presents the results of a survey, made during the summers of 1933 and 1934 of
the physiological behavior of the two species of Valonia which are most abundant at
Tortugas, Florida. One may well ask what justification there can be for yet another paper
on Valonia. Whatever the legitimate claims which may be made for such attention, they
are somewhat counterbalanced by the inaccessibility of Valonia, which has prevented that
examination by a variety of investigators which is the best safeguard against overemphasis.
Valonia macrophysa occurs at Tortugas only in the moat of old Fort Jefferson. This
organism demands complete protection from the effects of swell and surf. The growth
obtained on a horizontal ledge is luxuriant; that on an inclined or vertical surface sparse
and irregular. In the protected locations it demands, V. macrophysa is exposed to and
withstands, a wide range of light conditions and diurnal fluctuations in the composition of
the external medium. Valonia ventricosa is abundantly obtained on Bird Key Reef. The
distribution of V. ventricosa is complementary to that of V. macrophysa, and the solution
of the problem whether the species are distinct, raised thereby , must await adequate
transplant experiments. The range of sap composition which V. ventricosa and V.
macrophysa exhibit at Tortugas in sea water is described. Differences occur in the
composition of the sap of V. macrophysa grown in different parts of the moat of Fort
Jefferson. The principal causal factor appears to be the light condition which it obtains
during growth. In general the conditions which produce the most abundant growth of V.
macrophysa \ikewise produce the greatest concentration of potassium and lowest
concentration of sodium.
347. Stockard, C. R. 1908. Habits, reactions, and mating instincts of the "Walking Stick," Aplopus
348.
349.
mayeri. Papers Tortugas Laboratory 2: 43-59.
Carnegie Institution of Washington Publication Number 103.
This investigation of a protectively adapted insect is important to show definitely whether
the actions of such an animal are coordinated with its protective structure. It is concluded
that the habits of Aplopus mayeri on its food-plant Suriana maritima are as truly
protectively adapted as is its singular stick-like appearance.
. 1911. The influence of regenerating tissue on the animal body. Papers Tortugas
Laboratory 3, no. 41-48.
Carnegie Institution of Washington Publication Number 132.
It is stated that when the adult animal body begins to regenerate new tissue in order to
replace a lost part, or when abnormal secondary growths arise, the condition of growth-
equilibrium is disturbed and such a disturbance is followed by changes which affect the
usual physiological condition of the body. The question as to whether the changes
following normal regenerative growth are in any way similar to those effects resulting from
malignant or abnormal secondary growths arises.
. 1908. Studies of tissue growth. I. An experimental study of the rate of regeneration in
Cassiopea xamanchana (Bigelow). Papers Tortugas Laboratory 2: 61-102.
Carnegie Institution of Washington Publication Number 103.
The author responds to the studies of Zeleny (1903 and 1905) in which he suggested that
the greater the degree of injury within limits, the more rapid the rate of regeneration.
Zeleny suggested that the animal with the greater number of appendages removed might
exercise the regenerating ones more than the animal with less: activity should increase the
rate of regeneration in animals. The author tests the influence of rest and activity on
regenerating tissues of medusa and finds no increase in the regeneration rate from activity.
Rate of regeneration was also tested against food consumption, distance of cuts from the
margin of the medusa disks, cuts from different parts of variously shaped surfaces, removal
of oral epithelium of different sizes and at different distances, and the influences of
changed chemical conditions on regeneration.
87
350. Stoddart, D. R. and F. R. Fosberg. 1981. Topographic and floristic change, Dry Tortugas, Florida,
1904-1977. Atoll Research Bulletin 253: 1-56.
Topographic and floristic surveys of the Dry Tortugas Keys in 1904, 1915, and 1937 have
been used in discussions of the changing relationships between area and floristic diversity
on small islands over time, and of the processes of colonization and extinction. It is shown
that earlier topographic surveys are in general too unreliable to be so used. A list of Dry
Tortugas plants, including all published records was as well as new collections made in
1962 and 1977, is presented, together with maps of the keys made in 1977. The total flora
of about 130 species includes at least 35 native species, including 5 species of sea-grasses
and 4 species of mangroves. Introduced species are largely confined to the two largest
islands, and the floras of the smaller keys are dominated by a small number of native
species.
351. Stone, R. G. 1931-1932. Effect of irradiation by radium upon regeneration in marine annelids.
Carnegie Institution of Washington, Yearbook.
Note: published as follows: 1931, v. 30, p. 395; 1932, v. 31, p. 279.
The effect of combined beta and gamma radiations upon regeneration in polychaetes is
studied. Histological material is being used to determine the source of new tissue in
regenerated segments and to discover what tissues are affected by radiation.
352)
. 1934. Radium radiation effects on regeneration in Euratella chamberlin. Papers Tortugas
Laboratory 28: 157-66 (issued Jan. 1933).
Carnegie Institution of Washington Publication Number 435.
Regeneration in the polychaete annelids has been investigated in some instances but the
histological changes are not so well known as in the oligochaetes. The influence of X-rays
and radium upon regeneration in various animals has been demonstrated, but the
polycheates have seldom been used in these investigations. It has been found that the
effects of radiation are often limited to specific tissues; by reason of their greater
susceptibility they may be injured or destroyed by the exposure. During the summers of
1931 and 1932 the author was able to study the effects of radiation upon polychaete
regeneration at the Tortugas Laboratory. In the sabellid Euratella chamberlin, posterior
regeneration of abdominal segments is rapid and complete. Regeneration is inhibited by
sufficient exposure to the beta and gamma rays of radium. Similar exposure to gamma rays
alone has no effect upon the amount of regeneration. No structural changes were observed
in the radiated worms to account for this change. It is suggested that ionization induced by
the beta rays is responsible for the failure of regeneration .
393),
. 1936. Regeneration in the cirratulid Cirrineris. Papers Tortugas Laboratory 29: 1-12
(issued Nov. 1935).
Carnegie Institution of Washington Publication Number 452.
There has been considerable study of polychaete regeneration , but the observations are not
as extensive as those among the oligochaetes. This investigation of Cirrineris was
undertaken to determine the extent of segment replacement and the source of the new
tissues. Material was secured at the Tortugas Laboratory. In summary a head region and
six to seven segments posterior to it are regenerated when more than this number are
removed. Posterior regeneration is rapid and complete; the approximate number of
segments removed is replaced. Wound closure is effected in the same manner at both ends
of Cirrineris. The edges of the everted intestine unite with the epidermis to close the body
cavity. New nervous tissue arises by proliferation and inward migration of cells from the
adjacent epidermis. The old nerve cells do not participate in regeneration. Material for
regeneration of the intestinal lining arises by proliferation within the gut epithelium.
Mesodermal structures regenerate from old mesodermal tissues. Replacement material is
supplied by (a) nuclei and cytoplasm from muscles cells and connective-tissue elements
88
after degeneration of their differentiated cytoplasm; (b) peritoneal cells which furnish most
of the new material.
354. Stoneburner, D. L. and C. S. Harrison. 1981. Heavy metal residues in sooty tern, Sterna fuscata
tissues from the Gulf of Mexico and North Central Pacific Ocean. Science of the Total
Environment 19, no. 1: 51-58.
The comparison of mean cadmium, mercury and selenium concentrations in the eggs,
feathers and body tissues of breeding sooty tern (Sterna fuscata) from the Dry Tortugas,
Florida, and Lisianski Island, Hawaii, supports the hypothesis that a physiological
mechanism exists which functions in the detoxification of heavy metals. The data collected
from two geographically isolated populations of this pelagic bird indicate that the
mechanism responds in a uniform manner to widely different environmental levels of heavy
metals. The data and observations suggest that the mechanism evolved in response to
natural fluxes of heavy metal concentrations in the marine ecosystem, not in response to |
recent injections of heavy metal laden industrial wastes.
355. Stoneburner, D. L., P. C. Patty and William B. Robertson Jr. 1980. Evidence of heavy metal
accumulations in sooty terns. Science of the Total Environment 14, no. 2: 147-52.
Sooty terns from the population that nests at Bush Key, Dry Tortugas, Florida, had
substantial burdens of Cd, Hg and Se. Analysis of selected tissues, feces and eggs by
neutron activation techniques showed highest levels of Hg in eggs, feathers, and blood; of
Cd in kidney and bone; and of Se in kidney, liver, and feathers. The concentrations of Cd,
Hg, and Se in the eggs suggests that the heavy metals are being transmitted to succeeding
generations. The significance of the concentrations, their effect on the reproductive
success of the population, and the question of whether or not the metals transmitted to eggs
represent "bio-magnification" merit further work.
356. Strom, R. N.,R.S. Bramen , W. C. Jaap, P. Donan, K. B. Donnelly and D. F. Martin. 1992.
Analysis of selected trace metals and pesticides offshore of the Florida Keys. Florida
Scientist 55, no. 1: 1-13.
Trace metal and pesticide contents of sediments and producer and consumer organisms
were analyzed from samples taken from eighteen stations off the Florida Keys from
Biscayne National Park to the Dry Tortugas. Samples were analyzed for total mercury, tin
(inorganic and organic), arsenic (inorganic and methylated), lead, copper, cadmium, and
halogenated pesticides. Pesticide concentrations were below detection limits. In general,
concentrations of trace metals increased from sediments to producers to consumers at each
station. Though the concentrations tended to be low, some deviations were ascribed to
human inputs. Fewer significant correlations were observed than expected, possible
because of the dependence of the uptake mechanism upon the ability of the system
(sediment, producer, consumer) to remove trace metals from particular materials. Sponges
have this ability and may represent a useful means of monitoring the quality of the
environment on a sustained basis. The results are generally consistent with a relatively
clean environment with some localized anthropogenic effects.
357. Stromsten, F. A. 1911. A contribution to the anatomy and development of posterior lymph hearts of
turtles . Papers Tortugas Laboratory 3: 79-87
Carnegie Institution of Washington Publication Number 132.
This article concludes that the development of the posterior lymph hearts of turtles is
initiated by he vacuolation of the postiliac mesenchymal tissue during the middle and latter
part of the second week of development of the loggerhead turtle. The spongy tissue thus
formed is then invaded by capillaries from the first two or three dorsolateral branches of
the caudal portion of the postcardinal veins. The final stage in the development of the
posterior lymph hearts is reached by the dilation and confluence of these veno-lymphatic
358.
B59)
360.
361.
362.
363.
364.
89
sinuses, from before backward, forming a pair of sac-like organs, each with a single central
cavity.
——. 1910. The development of the posterior lymph hearts of the Loggerhead turtle
Thalassochelys caretta. Proceedings of the Iowa Academy of Science 17: 227-28.
Observations made on the lymphatic systems of turtles indicates their origin is more or less
independent of the venous system. Later investigations confirm this view, suggesting that
posterior lymph hearts of the loggerhead turtle are developed from embryonal cappillaries,
which have been captured and modified by the mesenchymal spaces of the post-iliac
regions of the body.
Tandy, G. 1931. The superficial structure of coral reefs; plant succession upon prepared substrata.
Carnegie Institution of Washington, Year Book 30, 32: 395; 26S.
Plant and animal successions were examined on concrete cubes planted in the water at
three sites: Fort Jefferson moat, an iron wreck east of Loggerhead Key, and northwest of
Loggerhead Key.
Tartar, V. 1938-1939. Regeneration in the starfish Linckia and in the protozoan Condylostoma.
Carnegie Institution of Washington, Year Book.
Note: published as follows: 1938, v. 37, p. 99-102; 1939, v. 38, p. 230-31.
Regeneration experiments were conducted on starfish with and without arms and isolated
arms. Under normal conditions polarity of arms is not altered by isolation. Tube feet cell
differentiation was examined in relation to color changes. In the ciliate, Condylostoma, the
normal form and typical arrangement of cytoplasmic differentiations may easily be altered.
Tashiro, S. 1914-1915. Further studies on CO) in sea water and CO) production in tropical marine
animals. Carnegie Institution of Washington, Year Book.
Note: published as follows: 1914, v. 13, p. 170; 1915, v. 14, p. 217-19.
Studies were conducted on the presence of "free CO 2 in sea water." A rapid method to
estimate amounts of CO, produced in sea water by marine animals was devised.
Taylor, J. B. 1981. Premetamorphic veligers of Fort Jefferson Dry Tortugas, Gulf of Mexico, and
Beaufort Inlet, North Carolina. Bulletin of the American Malacology Union, Inc. 50: 29-
30.
(No abstract available).
Taylor, W. R. 1928. The marine algae of Florida, with special reference to the Dry Tortugas.
Papers Tortugas Laboratory 25: 1-219.
Carnegie Institution of Washington Publication Number 379.
The study of marine vegetation of the Dry Tortugas was originally undertaken to provide a
simple check-list of algae of the islands for use of persons visiting the Carnegie Laboratory
there, with a description of the more important ecological features and records of the
locations where plants of experimental importance might be found. When it was discovered
that information about Florida algae in general was scanty, the study extended to a
thorough study of all available Florida material. Records of the occurrence of marine algae
on the east coast of Florida and the Florida Keys were collected. This is the first time, since
Harvey, Farlow and Melvill, that an attempt was made to list completely the Florida algae.
. 1925. The marine flora of the Dry Tortugas. Revue Algologique 2: 113-35.
The marine algae of the Dry Tortugas are listed, and a description of the distribution is
given of the important types throughout the area.
90
365. Teas, H. J. and P. B. Schroeder. 1971. Vegetation analysis in the Dry Tortugas by remote sensing.
Quarterly Journal of the Florida Academy of Science 34, no. (Suppl 1): 13.
Detailed ground truth observations were carried out on the four large islands in the Dry
Tortugas using aerial photography and 12S image enhancement equipment. Several
vegetation associations (strand-beach, strand-dune, strand-scrub) are distinguished and a
number of plant species identified (Rhizophora, Laguncularia, Bursera, Conocarpus,
Casuarina, Cocos, and Phoenix).
366. Tennent, D. H. 1911. Echinoderm hybridization. Papers Tortugas Laboratory 3: 117-51.
367.
368.
Carnegie Institution of Washington Publication Number 132.
The Toxopneustes female x Hipponoé male and the reciprocal cross Hipponoé female x
Toxopneustes male were easily made after allowing the eggs to stand in sea-water for some
hours before fertilization. In the embryos of both crosses made in ordinary sea-water,
which was alkaline, the Hipponoé influence showed a tendency to predominate. It is
suggested that the variations in the alkalinity of the sea-water, which have been brought
about artificially, may correspond to normal seasonal changes. The results of this and of
other investigations show species tendencies toward different grades of temperature and of
alkalinity. The explanation of the preponderance of one character over another in
echinoderm hybrids seems to lie in the reaction of the species toward a complex of factors.
. 1920. Evidence on the nature of nuclear activity. Proceedings of the National Academy of
Science 6: 217-21.
The author describes the results of the Arbacia eggs and other materials examined.
Basophilic bodies found are not in the nature of chromidia, but are the result of indirect
nuclear activity.
. 1942. The photodynamic action of dyes on the eggs of the sea urchin, Lytechinus
variegatus. Papers Tortugas Laboratory 35: 1-153.
Carnegie Institution of Washington Publication Number 539.
The work recorded in this paper was begun as a study of the experimental modification and
control of cell division in the egg of the sea urchin, Lytechinus variegatus. Early in the
investigation the photodynamic effects of the dye neutral red were found to be striking and
it was decided to undertake a study of the effects produced by other dyes. Transmission of
visible light by some of the filters was so low that the intensity of the light transmitted was
not sufficient to produce photodynamic effect. With dyes that produced a photodynamic
effect, irradiation of a solution of the dye resulted in the formation of a photocompound.
This photocompound was the active agent in the production of the photodynamic effect.
The threshold for violent surface reaction (blister cytolysis) of Lytechinus eggs in 1:150,000
solution of neutral red in sea water lay at about 2500 foot-candles. From this point to
about 4300 foot-candles violent surface reaction usually stood at about 2 per cent. Between
7000 and 7500 foot-candles it increased to 20-25 per cent, and between 800 and 9500 foot-
candles it increased to 75-90 per cent. At intensities from 300 to 10,000 foot-candles there
was a regular increase in the violence of the surface reaction and complete inhibition of the
cleavage processes. Irradiation in some of the solutions of dye at temperatures above 32° C.
resulted in injury from which the eggs did not recover. In blister cytolysis the formation of
blisters starts at a single point. Adjacent blisters come into contact with one another until
the entire surface is covered. The content of these blisters is liquid, is clear on the living
egg, and in the fixed egg seems to be the same as the cytoplasm of the egg with all formed
components removed. It is conceivable that these components could be filtered out, but
there is no evidence of the accumulation of granules at the point where the cytoplasm might
have been extruded .
9]
369. Tennent, D. H. and V. H. Keiller. 1911. The anatomy of Pentaceros reticulatus. Papers Tortugas
Laboratory 3: 113-16.
Carnegie Institution of Washington Publication Number 132.
This account is a description of the anatomy of Pentaceros reticulatus. Figures are used to
illustrate the organs which are described. Those which seem of greatest interest are the
intestinal caeca. These were found in some instances to be greatly distended, stimulation
causing their contraction. In this behavior we support the idea of the analogy of the
intestinal caeca of the starfish to the respiratory trees of the holothurian, an idea which has
been based upon the similarity of position of these organs.
370. Tennent, D.H., C. V. Taylor and D. M. Whitaker. 1929. An investigation on the organization in a
sea-urchin egg. Papers Tortugas Laboratory 26: 1-104.
Carnegie Institution of Washington Publication Number 391.
In this report the eggs of the sea urchin, Lytechinus, were studied from samples taken at the
Tortugas. The differentiation of ectoderm-forming substance over the entire surface of the
egg begins before fertilization by the exclusion of the endoderm-forming material from the
superficial layers of the egg. The number and relative distribution of micromeres is
independent of the plane of section and of the size of the fragment. There is no localization
of micromere-forming material.
371. Tennent, D. H., M.S. Gardiner and D. E. Smith. 1931. A cytological and biochemical study of the
ovaries of the sea-urchin Echinometra lacunter. Papers Tortugas Laboratory 27: 1-46.
Carnegie Institution of Washington Publication Number 413.
The investigations upon which this paper is based constitute a new method of attack on the
problem of the functional significance of chondriosomes, Golgi bodies and other
"inclusions" in protoplasm. In 1926 a definite research program for histochemical and
biochemical study of the eggs and ovaries of the sea-urchin Echinometra lacunter was
begun. Summarizing: analytical figures for the percentage of lipids and of glycogen are
given. The amount of the latter, 12.42 per cent, and 12.72 per cent of the dried extracted
tissue is high. In addition, the presence of cerebrosides and sphingomyelin are indicated.
The lipid composition of this tissue seems to be complex. The unsaturation of several
preparations used in the study of staining reactions was determined, to find out if there was
any correlation between unsaturation and staining with osmic acid (see Section I). On the
whole the lipids are probably more unsaturated than similar preparations from mammalian
tissues.
372. Thiele, J.. 1916. Molluskenfauna Westindiens. Zoologische Jahrbucher Supplement II: 109-32.
A listing of the mollusks of the West Indies is given and a preliminary catalogue of the
shell-bearing marine mollusks and brachiopods of the southeastern coast of the United
States. This article is in German.
373. Thompson, M. J. and T. W. Schmidt. 1977. Validation of the species/time random count technique
sampling fish assemblages at Dry Tortugas. Proceedings of the Third International Coral
Reef Symposium, No.1:283-288. Miami, Florida: RSMAS, University of Miami.
Ichthyofauna at four coral reef sites in Fort Jefferson National Monument, Dry Tortugas,
are compared during summers 1975 and 1976. Samples were taken using the species/time
random count technique, a newly developed visual censusing method based upon the rate at
which species are encountered by a free swimming observer. Data were collected by
different observers during the two years’ sampling. Within nine fish families dominating
the Tortugas ichthyofauna, the rank of five did not vary at all between 1975 and 1976
samplings. Among the four families exhibiting changes in abundance, only the Serranidae
showed a variation greater than 10.0%. The marked variation of 25.8% within this family
is attributed to identification problems within the genus Hypoplectrus. Overall numbers of
92
species and relative species abundances within each sampled coral reef area showed
minimal variation between years. The species rank correlation coefficient (Spearman's r s)
between two years of observations was 0.92. High correlation between results from two
different observation teams shows the species/time random count technique to be a highly
reliable method of comparing coral reef fish assemblages.
374. Thorp, E.M., A. Mann, T. W. Vaughan and F. J. Haight. 1936. Calcareous shallow-water marine
deposits of Florida and the Bahamas. With appendices: 1.Mann, A. Diatoms in bottom
deposits from the Bahamas and the Florida Keys; 2. Vaughn, Thomas Wayland. Current
measurements along the Florida Coral Reef Tract with notes on current observations,
Florida Keys, June, October, November, 1914. See separate entries for Appendices. Papers
Tortugas Laboratory 29: 37-143 (issued Dec. 1935).
Carnegie Institution of Washington Publication Number 452.
Determinations of the quantities of material derived from organic and inorganic sources
have yielded the following results: Coralline algae, collectively, are shown to be the
organic group that makes the largest contribution of organically secreted calcium
carbonate. Next in order of magnitude are the mollusks, followed in descending order by
foraminifera, madreporarian corals, alcyonarian spicules, worm tubes, crustacean
fragments, and Bryozoa. The principal non-calcareous mineral is quartz. Quantitative
counts of alcyonarian spicules show that they are relatively minor components of the
sediments, being exceeded by madreporarian fragments in a ratio of about 2.5 to 1.
Terrigenous minerals are remarkable scarce. A very small amount of volcanic glass and a
few species of heavy minerals occur well distributed over the region. The sources of all the
volcanic glass and some of the heavy minerals are thought to be distant and that they are
wind blown. Coal and ashes brought from outside sources by human agencies have been
introduced into the sediments of Tortugas lagoon, and, in smaller quantities, in a few other
places.
375. Tomas, C. R. and D. G. Baden. 1993. The influence of phosphorus source on the growth and
cellular toxin content of the benthic dinoflagellate Prorocentrum lima. Fifth International
Conference on Toxic Marine Phytoplankton, 565-70. St. Petersburg, Florida: Florida
Marine Research Institution.
The relationship between toxin content and nutritional status of the toxic marine
phytoplankton Prorocentrum lima was examined in a culture from the Dry Tortugas, grown
with inorganic phosphate and glycerol phosphate. Phosphorus source affected growth,
maximal densities, and okadaic acid content of Prorocentrum lima.
376. Torrey, H. B.. 1927-1928. Effect of thyroxin on division rates of various cells. Carnegie Institution
of Washington, Year Book.
Note: published as follows: 1927, v. 26, p. 228-229; 1928, v. 27 , p. 287.
Thyroxin depressed cell division and differentiation in eggs of sea-urchin (Echinometra
lacunter), ascidians (Phallusia nigra), and hydroids (Pennaria tiarella), collected at
Tortugas.
377. Treadwell, A. L. 1911. Eunicidae of Tortugas. Bulletin of the American Museum of Natural History
30: 1-12.
Systematic accounts of six species of polychaetous annelids are provided from specimens
collected in the dead coral rock around Fort Jefferson during 1908. Some species are
redescribed because of their earlier incomplete descriptions. Brief notes on their
abundance and distribution are included.
378. Treadwell, A. L. 1921. Leodicidae of the West Indian region. Papers Tortugas Laboratory 15: 1-
LS
93
Carnegie Institution of Washington Publication Number 293.
A systematic study based on specimens of the family Leodicidae is presented. Collections
were made at the Dry Tortugas and Key West region of Florida, and in Bermuda, Porto
Rico, Montego Bay, Jamaica, and Tobago. Collecting was done along shore or in
comparatively shallow water. The Leodicidae are a well-defined family in which the most
constant structures are internal rather than external. There is always a well-developed jaw
apparatus, composed of bilaterally arranged series of chitinous plates developed in a
pharyngeal pouch, and capable of protrusion for feeding purposes through the mouth. The
structure of these jaws was used by Ehlers as a basis for classification, though the external
organs are a more convenient means of recognition.
Se)
. 1917. Polychaetous annelids from Florida, Porto Rico, Bermuda, and the Bahamas. Papers
Tortugas Laboratory 11: 255-68.
Carnegie Institution of Washington Publication Number 251.
This paper is a preliminary description of some new species belonging to the Polychaetous
annelids, as well as new species of other families which have been collected incidentally in
this work, including a new sabellid belonging to the collection of the American Museum of
Natural History.
380. Ubelacker, J. M. 1982. Review of some little-known species of syllids (Annelida: Polychaeta)
described from the Gulf of Mexico and Caribbean by Hermann Augener in 1924.
Proceedings of the Biological Society of Washington 95, no. 3: 583-93.
The types of six little-known syllid species described by Augener in 1924 from the Dry
Tortugas, Florida, and from St. Thomas and St. Croix in the West Indies, were reexamined.
Haplosyllides floridana is a sexual form herein synonymized with it. Eusyllis antillensis
and Syllis (Typosyllis) tigrinoides are synonyms; the latter name is retained. Syllis
(Typosyllis) fuscosuturata has previously been synonymized with Branchiosyllis exilis
corallicoloides and remains a valid species.
381. Vaughan, T. W. 1914. The building of the Marquesas and Tortugas Atolls and a sketch of the
geologic history of the Florida Reef tract. Papers Tortugas Laboratory 5: 55-67.
Carnegie Institution of Washington Publication Number 182.
The study of the geology and the geologic processes of the Florida reef tract, and especially
of the Tortugas and the Marquesas, has been continued since 1910. It is now possible to
outline the salient geologic episodes in the history of the entire Florida Reef tract and to
Institution comparisons with other coral-reef areas.
382. ———.. 1910. A contribution to the geologic history of the Florida Plateau. Papers Tortugas
Laboratory 4: 99-185.
Carnegie Institution of Washington Publication Number 133.
This paper is the outgrowth of the author's association with two organizations, the United
States Geological Survey and the Carnegie Institution of Washington. The author has
visited all the principal keys between Miami and Key West, to collect and study bottom
samples, particularly the deposits accumulating behind the keys, and to examine several
important living coral reefs around the Tortugas. The scope of the paper was enlarged to
trace the geologic history of the Floridian Plateau from Oligocene to Recent time. This is to
be regarded as only a sketch of the geologic development of the Floridian Plateau, as many
problems need solution and many phases of its history need further investigation. Perhaps
its principal value may be in directing attention to some of the unsolved problems. It is
necessary to know more accurately the amount of water discharged by the streams and the
quantities of solids borne by them to the sea. The chemical processes of precipitation have
not been sufficiently studied. There is also great need for more extensive studies of the
marine bottom deposits within the 100-fathom curve. The deep wells recently put down on
94
383.
384.
S85:
Key Vaca, Big Pine Key, and Key West have given valuable data, but deep wells are also
needed on the Marquesas, and the Tortugas, in order to discover what underlies the surface
formations. It is hoped this paper may serve as a convenient summary of the present
knowledge of the geologic history of this interesting region, perhaps present an
interpretation somewhat different from those preceding, and be a stimulus to further
investigation.
. 1915. Coral-reefs and reef corals of the southeastern United States; their geologic history
and significance. Bulletin of the Geological Society of America 26: 58-60.
The geologic history of the extensive coral reefs of the southeastern United States and
near-by West Indian Islands was outlined, and the bearing they have on the theory of coral
reef formation was indicated. The author stated his conclusions regarding the Florida coral
reefs as follows: (1) Corals have played a subordinate part, usually a negligible part in the
building of the Floridean plateau; (2) every conspicuous development of coral reefs or
reef corals took place during subsidence; (3) in every instance the coral reefs or reef corals
have developed on platform basements which owe their origin to geologic agencies other
than those dependent on the presence of corals. The conclusions in this report are
summarized as follows: Critical investigations of corals as constructional geologic agents
are bringing increasing proof that they are not as important as was believed. All known
modern offshore reefs which have been investigated grow on platforms which have been
submerged in Recent geologic times. No evidence has been presented to show that any
barrier reef began to form as a fringing reef and was converted into a barrier by subsidence.
There were platforms in early Teritiary time on the site of many of the present-day
platforms, and evidence has yet been adduced to prove long-continued, uninterrupted
subsidence in any coral-reef area. The width of a submerged platform bordering a land
area is indicative of the stage attained by planation movement. The importance of coral
reef studies to geology suggests they are only a conspicuous incident in time.
. 1936. Current measurements along the Florida coral reef tract. Papers Tortugas Laboratory
29: 129-41.
Carnegie Institution of Washington Publication Number 451. Note: This is Appendix 2 to
Calcareous shallow water marine deposits of Florida and the Bahamas by Eldon Marion
Thorp.
During June and July 1914, while studying the phenomena associated with the Florida
Coral-Reef Tract the author initiated a series of current measurements by using Ekman
current meters. South of the Tortugas a non-tidal current toward the west is clearly
indicated. The data here presented are inadequate for positive conclusions regarding the
Counter Current.
. 1915. The geologic significance of the growth rate of the Floridian and Bahamian shoal-
water corals. Journal of the Washington Academy of Science 5, no. 17: 591-600.
The object of this investigation has been to aid in understanding the amount of work stony
corals may do as constructional geologic agents, and especially in the formation of coral
reefs. This subject needs to be studied from at least five different view points , e.g.: (1) the
quantity of material contributed by corals and that contributed by other agents must be
estimated and the respective proportions determined; (2) in coral reef areas the ratio of the
area covered by corals to that not covered by corals should be estimated; (3) the relations
of coral reefs, continuity and discontinuity must be determined; (4) marine bottom deposits
must be analyzed according to the source of the material, and the percentage of the calcium
carbonate contributed by the differing agents estimated; (5) the rate of growth of corals
needs to be known. There is no single formula for the growth rate of corals, as it varies by
species and ecologic conditions. Observations/experiments on the growth rates of Tortugas
corals are as follows: (1) Colonies obtained from the planule whose history was known,
386.
387.
388.
95
and were planted off the moat wall and on the NW side of Loggerhead Key; (2) Colonies
cemented to tiles and planted at the same sites as above; Colonies naturally attached at sites
described above. The reef species of greatest concern and importance is Orbicella
(Montastrea) annularis followed by in importance, Maeandra strigosa, M.
labyrinthiformes, and Siderastrea siderea. The upward growth is critical of the massive
heads Orbicella (Montastrea) annularis which form the strong framework of the reef and
averages | foot in 43.54 years or 7 mm /year and which might form a reef 150 feet thick in
between 6500 and 7600 years. A table on the average annual growth rates of corals from
the Florida region is provided.
. 1914. The platforms of barrier coral reefs. Bulletin of the American Geographical Society
46: 426-29.
The author states that there are three kinds of coral reefs: fringing or shore reefs which
occur along the strand line, barrier reefs which occur at varied distances off shore and have
lagoons from one to as much as forty fathoms depth between them and the strand line, and
atolls, which are ring-like and enclose lagoons. As the relations of barrier reefs and atolls
to the platforms on which they stand constitute the essential part of the theory of
development of Recent reefs, the discussion of coral reef theory has been waged over the
interpretation of these relations. The object of this paper is to point out the relations of
barrier coral reefs to the last dominant change in position of the strand line and to indicate
the organisms forming Recent barrier reefs have played in building the reef platforms.
. 1914. Preliminary remarks on the geology of the Bahamas, with special reference to the
origin of the Bahamian and Floridian Oolites. Papers Tortugas Laboratory 5: 47-54.
Carnegie Institution of Washington Publication Number 182.
The author presents a preliminary summary of the information compiled (to 1914), on the
origin of calcium carbonate sediments in south Florida, and the Bahamas, using various
hypotheses developed by the leading geologists of the time. These geologists included
Alexander and Louis Agassiz, Sanford, and Drew who worked in the Dry Tortugas and
believed the precipitation of calcium carbonate was due to the effects of denitrifying
bacteria.
. 1916. The results of investigations of the ecology of the Floridian and Bahamian
shoalwater corals. Proceedings of the National Academy of Sciences 2: 95-100.
This paper presents a summary of the knowledge on the ecology of shallow water corals in
the Florida-Bahamian region, with a detailed description of new information on food
preferences of corals, and salinity and water temperature tolerances, based on studies
conducted at the Dry Tortugas and the upper Florida Keys area. Mayer, at the Tortugas
Laboratory, found that temperatures of 13.9 °C would exterminate the principal Floridean
corals; similar results were found for corals around Australia. Light experiments at Fort
Jefferson suggested vigorous coral growth in well lit wharf areas, and little growth in piling
areas of perpetual shading. Tests conducted at the Marine Laboratory suggested that corals
could survive at salinities of 27-38 ppt, but not as low as 19 ppt. Other conditions
necessary for vigorous coral growth are maximum water depths of 45 meters and rocky or
firm bottoms, without silty deposits. The growth rate of corals was determined by planting
planulae in the laboratory, by measuring colonies which had been cemented to disks, and
fixed on heads of stakes driven into the sea bottom. Measurements of colonies naturally
attached were also made. Plantings at the Tortugas were made at Loggerhead Key, and
around Fort Jefferson. The more massive the coral, the slower the growth; while the more
ramose (Acropora palmata) and the more porous the skeleton, the more rapid the growth.
The growth rate of the principal reef builders (massive corals) in the Florida region,
Orbicella (Montastrea) annularis is from 5-7 mm per year and would form a reef of 150
96
389.
390.
391.
feet in thickness from 7620 to 6531 years. A. palmata may build a similar thickness in
1800 years.
. 1914. Sketch of the geologic history of the Florida coral reef tract and comparisons with
other coral reef areas. Journal of the Washington Academy of Sciences 4: 26-34.
The author presents the two hypotheses for the formation of atolls: one attributes atolls to
the submarine solution of the interior of a mass of limestone; the other accounts for them
by constructional agencies. The author believes the solution theory is disproved by a
chemical examination of sea-water from a Tortugas lagoon. He believes the Marquesas
and the Tortugas are constructional phenomena and owe their configuration to the
prevailing winds and currents.
. 1918. The temperature of the Florida Coral-Reef Tract. Papers Tortugas Laboratory 9:
319-39.
Carnegie Institution of Washington Publication Number 213.
The temperature data presented were assembled primarily for their bearing on the effect
temperature exerts on the bathymetric and geographic distribution of coral reefs.
Temperature is also one of the most important factors in determining the geographic
distribution of sea-level and near sea-level reefs.
Vaughan, T. W., M. A. Goldman, J. A. Cushman, M. A. Howe and others. 1918. Some shoal-water
bottom samples from Murray Island, Australia, and comparisons of them with samples
from Florida and the Bahamas. Papers Tortugas Laboratory 9: 235-97.
Carnegie Institution of Washington Publication Number 213.
The present paper is a preliminary contribution to the study of the marine bottom deposits
in three coral-reef areas: Murray Island, Australia; the Bahamas, and southern Florida.
Mechanical analyses have been made of all samples except those obtained in 1915, and the
results of the chemical analyses of a selected set are presented. An attempt has been made
to outline a method of studying calcium carbonate bottom deposits, in the hope that
progress may be made toward an adequate classification of such sediments. The Tortugas
Lagoon samples are coarser than those in Marquesas Lagoon, and those from the latter
locality are coarser than the Bahama sample from South Bight and the west side of Andros
Island. Some terrigenous material, mostly quartz sand, is washed into Biscayne Bay,
Florida, and into the sounds south of it, but otherwise practically none reaches the key and
reef region. The Florida area is therefore a perfect example of limestones forming in shoal
water near a land area which is not crossed by large streams. The Fe,0; content of the
Florida samples seems somewhat higher, up to about 0.37 per cent, than that of the Bahama
samples. Reconsideration of the evidence bearing upon the precipitation of CaCo3 in
tropical and subtropical waters and the possibility of its re-solution by ocean-water leads to
the conclusion that precipitation is resulting from both organic and inorganic agencies, and
that no appreciable re-solution is taking place in the water.
392. Visscher, J. P. 1930-1931. Distribution of barnacles with special reference to behavior of larvae.
Carnegie Institution of Washington, Year Book
Note: published as follows: 1930, v. 29, p. 346; 1931, v. 30, p. 397.
A study of the barnacles was made, more than twenty species being found. Several species
appear to be new to science. Behavior of larvae appeared to vary depending on habitat, as
certain barnacles were found on crabs, others above the tide on pilings, still others only on
coral and on the spiny lobster, Panulirus argus.
393. Vukovich, F. M. 1988. On the formation of elongated cold perturbations off the Dry Tortugas.
Journal of Physical Oceanography 18, no. 7: 1051-59.
The life cycle of a cold perturbation on the boundary of the Loop Current in the Gulf of
:
97
Mexico was studied over the period of 18 March to 22 May 1984, approximately a 60-day
period. The study focused on the behavior of the surface and subsurface area of the cold
perturbation as it moved along the boundary of the Loop Current. The area of the
perturbation was defined by an alongflow-scale length, which is the scale length parallel to
the unperturbed flow of the Loop Current, and the crossflow-scale length, which is the
scale length perpendicular to the unperturbed flow of the Loop Current.
394. Vukovich, F. M. and G. A. Maul. 1985. Cyclonic eddies in the eastern Gulf of Mexico. Journal of
Physical Oceanography 15, no. 1: 105-17.
Cold-domed cyclonic eddies juxtaposed to the cylconic shear side of the Gulf Loop
Current are observed in simultaneously obtained hydrographic, current meter mooring, and
satellite data as cold perturbations on the northern extreme of the current and grow either
into a cold tongue or a quasi-stable meander off the Dry Tortugas, Florida. Areal
shipboard surveys show closed isopleths of temperature and salinity, and surface
geostrophic current speeds relative to 1000 db are in excess of 100 cms super(-1). The
diameter of the cold domes varied from 80 to 120 km.
395. Wallace, W. S. 1908. A collection of hydroids made at the Tortugas during 1908. Carnegie
Institution of Washington, Year Book 7: 136-37.
At least fifty species of hydroids were collected. A tentative list of those identified is
provided.
396. Wartman, W. 1929. Studies on Echinometra. Carnegie Institution of Washington, Year Book 28:
Die
(No data report provided).
397. Watson, J. B. 1908. The behavior of noddy and sooty terns. Papers Tortugas Laboratory 2: 187-
Daas
Carnegie Institution of Washington Publication Number 103.
The work presented in this report is preliminary in nature. Following a general description
of the two species, a geographical situation and present history of the tern colony at the
Tortugas is given. All observations were recorded during the nesting season on Bird Key,
a small coral island covered in part by bay cedar, mixed with cactus in the central western
parts of the island, with little vegetation elsewhere. Observations on their foods and
feeding habits indicated that the birds usually feed in groups, never swim nor dive, but
skim along the surface picking up small fish being attacked by larger fish. Feeding
distance from the Key was estimated between 4 and 10 knots. Mating has been suggested
prior to arriving at the Tortugas, although some indications of sexual activity occurred for
the noddies, but not the sooties. Noddies nest in vegetation, while sooties build nests in
sand. Usually one, sometimes two eggs are laid, with a period of incubation for the noddy
from 32 to 35 days. The parents alternately feed the young at intervals from 1 to 4 hours;
their general conduct does not greatly change at the arrival time of the young. However,
two days after the arrival, the parents are more ferocious; both species return to normalcy
as the chicks gain strength. The birds become exhausted caring for their young, and collect
upon the beach for "sunning". Egg coloration tests indicated that neither species
recognized its own egg. As for the noddy's nest environment, it could be disturbed without
affecting the bird, as long as the egg position was not changed. Tests conducted using
Porter's learning maze indicated that noddy's were slower than sooties, because of their
longer standing time. Further maze tests using darkness and maze rotation were
inconclusive. Other in captivity tests showed that the sooty is highly excitable and nervous,
whereas the noddy is stolid and indifferent.
98
398. Watson, J. B. and K. S. Lashley. 1915. Homing and related activities of birds. Papers Tortugas
Laboratory 7: 1-104.
Carnegie Institution of Washington Publication Number 211.
The present series of studies on the behavior of birds is a direct outgrowth of an
investigation made on the noddy and sooty terns nesting on Bird Key, Tortugas, Florida.
The homing "instinct" is the central topic in all the papers. In the 1907 investigation the
fact appeared that terns possess a homing sense, behaving exactly as do homing pigeons
when sent away from their nests and young. The 1907 investigation already referred to is
concerned largely with instincts in terns-those of feeding, nesting, brooding, etc. In general
the problems of proximate orientation are relatively simple and straightforward. On the
island of Bird Key the terns make their adjustment to the nest, mate, young, etc., on the
basis largely of visual habits. There is no evidence of any remarkable or unusual
sensitivity, nor of the functioning of any hypothetical sense-organ. The present paper
seems to call for a separation between proximate orientation and distant orientation.
Mathematical considerations show that at such distances the goal can not possibly
(directly) visually stimulate the bird, even granting absolute visual acuity and complete
absence of haze, etc. This work has shown further, in the terns at least, that there is no
special Spiirsinn-special tactual or olfactory mechanism situated in the nasal cavity which
may function in homing. The task of explaining distant orientation is an experimental one,
which must yield positive results as soon as proper methods are at hand. Two lines of
investigation offer hopeful results: the rearing of homing pigeons in a cote, or the rearing
of the birds in a wire-covered yard attached to a cote. We could tether individual birds to
the top of the cote by cords which would permit a view only of the neighborhood
immediately surrounding the cote. With these experiments upon homing, work upon the
sensory equipment of the homing pigeon should be carried on. It is just possible that these
animals possess on certain parts of the body, tactual and thermal mechanisms which may
assist them in reacting to slight differences in pressure, temperature, and humidity of air
columns. The experiments and conclusions on homing proper can be found on pages 59
and 60. These results, which not settling the question of the sensory mechanism by means
of which the birds return to the nests, do remove all doubts about the fact that the noddy
and sooty terns can return from distances up to 1,000 miles in the absence of all landmarks.
The problem of homing has thus become defined, and experimental work of a definite kind
is needed for its solution .
399. Wells, R. C. 1922. Carbon-dioxide content of sea water at Tortugas. Papers Tortugas Laboratory
18: 87-93.
Carnegie Institution of Washington Publication Number 312.
It is generally considered that the carbon-dioxide content of sea-water may be increased by
accessions from the air, by animal life, by the decay of organic matter in the sediments on
the bottom or elsewhere, by the solution of carbonate rocks, by the contributions of rivers,
and by gas vents beneath the sea. Sea-water may lose carbon dioxide to the air, to plants,
and in the formation of carbonate rocks and the carbonaceous parts of organisms. Mere
evaporation and precipitation also alter the carbon-dioxide concentration somewhat if other
conditions remain unchanged. The writer made determinations on sea-water from
Tortugas, Florida, in June 1919 taken directly from the sea at various points about
Loggerhead Key, which reveal unmistakable diurnal variations. The water has sufficient
contact with plants and sea-weeds to show the effect of photosynthesis on its CO, content.
There is a loss of CO, by day and a gain by night. Plant life appears to be the chief agency
in causing a daily variation in the CO, content. Determinations of CO, should probably be
made soon after the time the samples are collected, on account of the possibility of the
decay of organic matter, such as algae, in preserved samples. The average "excess base"
found at Tortugas corresponds to a normality of 0.00239. This titration includes
everything that consumes acid; it represents chiefly bicarbonate, about 0.00183, some
99
carbonate, about 0.00041, and other substances that contribute to the alkalinity, about
0.00015. The methods used in arriving at these figures were provided along with a record
of determinations made at Tortugas and the relation between the carbon-dioxide content of
the water and time of day.
400. Westinga, E. and P. C. Hoetjes. 1981. The intrasponge fauna of Spheciospongia vesparia (Porifera,
Demospongiae) at Curacao and Bonaire. Marine Biology 62, no. (2-3): 139-50.
The infauna of 35 individuals of Spheciospongia vesparia (Lamarck, 1814) of different
volumes and from different sites and depths have been inventoried and compared. The
number of sponge-inhabiting taxa is logarithmically related to sponge volume. Biomass
and total number of the animals contained in the sponge are directly proportional to sponge
volume. Numerical and taxonomic composition of infaunas from different sampling sites is
fairly constant. Biomass and total number of sponge inhabiting animals is not significantly
different for any of the four sampling sites. Several taxa, however, are more abundant in
sponges from one or more localities. The ratio of total biomass to total number of
intrasponge fauna is found to be significantly smaller for sponges collected in deep water
than in shallow water. Differences from and similarities with Pearse's results (1932,1950)
on the infauna of the same sponge species at Dry Tortugas and Bimini are discussed. The
relation of the number of contained taxa and the volume of a sponge is compared with the
relation of island size and number of taxa present according to MacArthur and Wilson's
island theory (MacArthur, 1972). Finally the erratic occurrence of some taxa as opposed to
the highly regular occurrence of some other taxa is discussed. It is concluded that the
composition of the sponge-infauna in specimens larger than 11 is highly constant and that
the sponge-inhabiting fauna constitutes an ecological community.
401. Westrum, B. L. and P. A. Meyers. 1978. Organic carbon content of seawater from over three
Caribbean reefs. Bulletin of Marine Science 28, no. 1: 153-58.
Seawater samples from transects crossing three Caribbean coral reefs, including the Dry
Tortugas, showed variations in concentrations of organic carbon. Total organic carbon
increased substantially over two fringing reef crests. Most of this increase occurred in the
particulate fraction at the seaward edge of the crest but in the dissolved fraction at the
landward edge. Back reef levels of total organic carbon were lower than those seaward of
the reef. These observations support the hypothesis that organic carbon can be physically
removed from the benthos at the turbulent reef crest and be subsequently utilized in
backreef areas. The reef-flat formation studied in January 1975 in the Dry Tortugas was
situated off the western shore of Loggerhead Key. This study indicates that organic matter
contributed at the crest is available as a resource to only a limited portion of the backreef
community - that part located directly behind the crest. The observed decrease in TOC
levels implies quick biological utilization or loss through physical processes. Thus, despite
continual input, no net accumulation of organic matter occurs in the backreef area, and this
region can be described as being relatively depleted in organic carbon. If large coral
formations are present, as at the Dry Tortugas location, they can contribute organic matter
to the surrounding seawater.
402. Wheaton, J. 1980. Ecology of gorgonians (Octocorallia: Gorgonacea) at Dry Tortugas, Florida .
Florida Scientist, 43 (suppl. 1), 20.
This study reports the species composition and distribution of the gorgonian fauna of Long
Key Reef, Dry Tortugas during the summers of 1975-1976. 23 species were recorded.
Additional samples increased the number of species to 35. Most shallow reef gorgonians
were Plexaura.
403. Wheaton, J.L., W.C. Jaap, B. L. Kojis, G. P. Schmahl, D. L. Ballantine and J. E. McKenna Jr.
1993. Transplanting organisms on a damaged reef at Pulaski Shoal, Ft. Jefferson National
100
Monument, Dry Tortugas, Florida, USA: An experiment to enhance recruitment. (abs.).
Proceedings of the Seventh International Coral Reef Symposium, p. 639. Mangilao, Guam:
University of Guam.
Grounding of the 475-ft. freighter, Mavro Vetranic, at Pulaski Shoal Reef, Dry Tortugas,
on 30 October 1989, damaged 3,465 m’ of reef surface. After one year, minimal
recruitment of macrobenthos, principally the alga Dictyota, had occurred. An experiment
was designed to test effects of adding relief and transplanting sponges, octocorals, and
scleractinian corals on recruitment of biota to the damaged area. In Sept. 1991, one control
and two experimental sites, each 9-m’, were selected, marked, mapped, and photographed.
Large reef rocks were placed in one experimental plot to provide relief and refuge. More
than 185 organisms (73 species of algae, Porifera, and Cnidaria) were transplanted and
cemented into five of the nine square-meter subunits in the other experimental plot.
Transplanting was labor intensive, requiring 64 man-hours to collect, move, and cement
organisms. We then rephotographed and mapped the plots. Sites will be monitored to
determine if recruitment of macrobenthic organisms is enhanced .
404. Whitaker, D. 1926. Organization of echinoderm egg, and a measurable potential difference between
the cell interior and outside medium. Carnegie Institution of Washington, Year Book 25:
248-55.
Egg development, investigated in the sea-urchin, Lytechinus, suggested that the
differentiation of ectoderm begins before fertilization by the exclusion of the endoderm-
forming substances from the superficial layers of the egg. Micromere-forming substances
do not differentiate before fertilization.
405. White, S.C., W. B. Robertson Jr. and R. E. Ricklefs. 1976. The effect of Hurricane Agnes on
growth and survival of tern chicks in Florida. Bird-Banding 47, no. 1: 54-71.
In June, 1972 Ricklefs and White were studying the energetics of nestling growth in sooty
terns (Sterna fuscata) at the Dry Tortugas, when Hurricane Agnes passed west of the area.
High winds, heavy rain, rough seas, and low temperatures prevailed for more than a week.
Robertson worked in the colony from 28 June, about a week after the storm subsided, to 6
July. It is reported here the effects of Hurricane Agnes on the growth and survival of
young sooty terns and brown noddies (Anous stolidus ).
406. Wichterman, R. 1942. Cytological studies on the structure and division of three new ciliates from
the littoral earthworm of Tortugas. Papers Tortugas Laboratory 33: 83-103 .
Carnegie Institution of Washington Publication Number 524.
During the summer of 1939 this study of Protozoa inhabiting the intestine of the littoral
earthworm Pontodrellus bermudensis Beddard, was begun. The study revealed three
previously undescribed ciliates: Hysterocinita pontodrila, n. s.p.; Anoplophyra
macroneucleata, n. sp.; and Maupasella leptas, n. sp. This paper describes the ciliates and
gives an account of fission in each species. Of the 230 worms examined, 64% were
infected with the ciliates. Generally a worm was parasitized with two different species.
Observations on the length of life of the ciliates in seawater were recorded. Encystment
was not encountered. The presence of stages in the life history of acephaline gregarines
and nematodes was noted.
407.
. 1942. A new ciliate from a coral of Tortugas and its symbiotic zooxanthellae. Papers
Tortugas Laboratory 33: 105-11.
Carnegie Institution of Washington Publication Number 524.
A new ciliate was found on the coral Eunicia crassa E. and H. and is described as
Paraeuplotes tortugenesis, n. gen. and n. sp., and is placed in the family Paraeuplotidae, n.
fam. The coral it was found on is a member of the Alcyonaria fauna, and is commonly
found in the Caribbean, and is abundant on the reefs of the Tortugas. The morphology of
101
the ciliate is discussed, as well as the presence of zoozanthellae. The question 1s posed
"what is the nature of the symbiosis between the protozoan and the zooanthellae it
contains?".
408. Williams, O. L. 1932. Studies on the nematodes of Tortugas fishes. Carnegie Institution of
Washington, Year Book 31: 291-92.
Observations of more than 800 fishes representing about 175 species during the summer of
1932 demonstrate that the incidence of infestation with nematodes is lower in fishes of the
Tortugas than in the cooler, shallow waters found farther north.
409. Willier, B. H. 1936. A study of the early embryology of the Loggerhead sea turtle and of sharks.
Carnegie Institution of Washington, Year Book 35: 92.
The embryological development of the Loggerhead turtle (Caretta caretta) was examined
from the time of egg laying to within a few days of hatching. Significant observations are
presented.
410. Wilson, C. B. 1936. Parasitic copepods from the Dry Tortugas. Papers Tortugas Laboratory 29:
327-47 (issued Dec. 1935).
Carnegie Institution of Washington Publication Number 452.
Two collections of parasitic copepods contained in the present paper were made at the
Marine Laboratory of the Carnegie Institution in the Dry Tortugas, involving the handling
of a large number of the local fishes. In addition to the specific objects of investigation it
was soon noted that the fish were more or less infested with parasitic copepods and
isopods. Upon identification, seven of the species are new to science, and two others have
been made the types of new genera. The other species have been obtained before either in
the waters around the Dry Tortugas, the Bahamas, the Bermudas or the West Indies.
411. Winegarner, C. E., W. B. Robertson and W. Hoffman. 1984. Anolis sageri sageri (brown anole)
USA: Florida: Monroe Co: Dry Tortugas, Garden Key. Herpetological Review 15, no. 3:
77-78.
Three males and one female specimen were taken on a large pile of bricks and rubble just
east of the moat surrounding Ft. Jefferson, April 8-10 1983. Population currently seems
limited to this small portion of the island, so introduction may have been very recent. A
construction barge moored adjacent to the collection site from October 1981 to June 1982
possibly was a source of colonizing individuals. However the regular arrival of Park
Service boats and private vessels are other possibilities.
412. Wolfe, C. A. 1989. "Growth of the Brown Noddy (Anous stolidus) in the Dry Tortugas (Florida)."
Master of Science, Florida Atlantic University, Boca Raton.
The author discusses the slow growth rate of the brown noddy nestlings in the Dry
Tortugas as to what would be predicted based on adult body size and mode of
development. This prolonged growth pattern is typical of tropical pelagic seabirds. An
intraspecific comparison of growth rates among several populations of brown noddies ,
indicates that growth of body mass of the Tortugas noddies is significantly faster, the
development period shorter, and the asymptotic size smaller than in Pacific populations.
However, there were no differences among the populations in the rates of wing or culmen
growth. The Bush Key nestlings appear to receive a higher quality diet that contains
proportionally more fish, while Pacific nestlings receive substantial amounts of squid. The
Pacific nestlings also seem to be subjected to a thermally more stressful microclimate,
which may necessitate the allocation of proportionally more of their total energy to
thermoregulation and less to growth.
102
413.
414.
415.
Woolfenden, G. E. and W. B. Robertson Jr. 1991. A banded red knot seen at the Dry Tortugas.
Florida Field Naturalist 19, no. 4: 106-7.
The red knot (Calidris canutus) is a locally abundant and winter visitor on both coasts of
Florida, however it is rare at the Dry Tortugas, with only three sightings prior to a sighting
made by the authors during May-June 1988. It was suggested that all sightings at the
Tortugas represented birds of a knot population that winters along the Atlantic coast of
Patagonia in South America.
. 1975. Least terns nest at the Dry Tortugas. Florida Field Naturalist 2, no. 1: 19-20.
On July 1, 1973, as members of the tern-banding party landed on Middle Key, Dry
Tortugas, they saw 4 adult-plumaged Least terns and one fledged juvenile in the company
of 8 adult-plumaged Roseate Terns, Sterna dougallii. Search of the island, a barren sand
bank with only a small area above high tide, revealed 2 Least Tern nests, one with 2 eggs
and the other with one egg, and 4 Roseate nests, each with 2 eggs. It is of interest that Least
Terns have again attempted to nest at Dry Tortugas after an absence of almost 25 years.
Woolfenden, G. E., S.C. White, R. L. Mumme and W. B. Robertson Jr. 1976. Aggression among
starving cattle egrets. Bird-Banding 47, no. 1: 48-53.
Cattle egrets (Bubulcus ibis) flying over the Gulf of Mexico often land at the Dry Tortugas.
Food suitable for cattle egrets is scarce locally and many egrets die at the Dry Tortugas,
presumably from starvation. In June 1975 an infestation of a sea grape tree by caterpillars
of the moth Sarasota Plumigerella Hulst. provided a natural, albeit limited, food supply at
which we observed cattle egret behavior. From several observers, we were able to
compare aggression, feeding frequency, plumage condition and death weights of the
starving birds. Of special interest was the opportunity to test relationships between
aggressiveness and feeding frequency under the unusual circumstances of starving birds
competing for a concentrated, but limited food supply.
416. Woolfenden, G. E and W. B. Robertson Jr. 1975. First nesting of the house sparrow at Dry
Tortugas. Florida Field Naturalist 3: 23-24.
This paper describes the first occurrence of the House sparrow at the Dry Tortugas.
During mid-June 1974, nest building and copulation was observed in a coconut palm east
of the moat bridge on Garden Key. Four eggs were found later, but no further inspections
were made. House sparrows that reach the Tortugas are considered true migrants or birds
from the West Indies that accompanied north-bound migrants of other species.
417. Yamanouchi, S. 1929-1935. Life histories and cytology of marine algae. Carnegie Institution of
Washington, Year Book .
Note: published as follows: 1929, v. 28, p. 297; 1930, v. 29, p. 346; 1931, v. 30, p. 371;
1932, v. 31, p. 259; 1933, v. 32, p. 265; 1934, v. 33, p. 263; 1935, v. 34, p. 75.
Local populations of marine algae, Phaeophycae and Chlorophyceae were studied with
emphasis on Caulerpa. Reproductive phases of many specimens were collected for later
morphological and cytological study.
418. Yonge, C. M. 1936. Studies on the biology of Tortugas corals. I. Observations on Maenadra
areolata Linn. Papers Tortugas Laboratory 29: 185-98 (issued Dec. 1935).
Carnegie Institution of Washington Publication Number 452.
Maenadra areolata is one of the commonest corals of the Caribbean and Florida reefs. It
is a highly specialized species adapted for life in a restricted environment. It thrives best
on the flats behind the reefs. It has no firm basal attachment, it can not resist the impact of
the waves of rough seas. At the Tortugas it is very common in sheltered areas on the inner
side of the circle of reefs where wide stretches of sand occur. The best collecting ground is
the lee of Bird Key Reef. Feeding is entirely by means of the tentacles, there is no reversal
103
of ciliary currents. Not only is sediment removed very rapidly from the surface, but
colonies can completely uncover themselves within twelve hours after being buried in the
sad. Unlike Fungia which uncovers itself by the exclusive action of cilia, M. areolata first
distends the tissues with water. Distension for cleansing is essentially different from
expansion for feeding. Planulation, so far as can be determined at present, has a lunar
rhythm, culminating about the time of new moon. After an initial stage when upward and
outward growth are about equal, outward growth predominates, an oval or rounded colony
with a small basal attachment being finally produced. The stage at which detachment
occurs must vary with environmental conditions. Colonies may be formed from a single
planula or from the fusion of several. M. areolata is a species highly adapted for life on
sand occupying in the Atlantic, the same habitat occupied by the Fungiidae in the Indo-
Pacific. Adaptability in reef-building corals is discussed.
419.
. 1936. Studies on the biology of Tortugas corals. IJ. Variation in the genus Siderastrea.
Papers Tortugas Laboratory 29: 199-208 (issued Dec. 1935).
Carnegie Institution of Washington Publication Number 452.
This paper presents data on stony corals on the Tortugas reefs. The genus Siderastrea is
represented at the Tortugas by the two species, S. siderea and S. radians. S. siderea forms
larger rounded colonies which were not observed above the level of low-water springs. S.
radians is essentially a shore-living species possessing the physiological adaptations
characteristic of all shore-living animals. S. radians is capable of great modification both
in the form of the skeleton as a whole and also in the size and shape of the corallites and in
the number, slope and thickness of the septa. This species has been enabled, as a result, to
occupy a variety of habitats, the extremes being represented by the surf region on the beach
rock on the one hand, and by the still, sediment-laden water in the moat at Fort Jefferson on
the other. The relation between form and environment in corals is discussed and the
general conclusion reached that the great success of the Madreporaria is probably due to
the presence of species highly specialized for a particular environment and also of others
which can be modified for existence in a variety of different environments .
420. . 1937. Studies on the biology of Tortugas Corals. III. The effect of mucus on oxygen
consumption. Papers Tortugas Laboratory 31: 207-14 (issued Oct. 1937).
Carnegie Institution of Washington Publication Number 475.
Experiments are described which indicate that a large proportion of the apparent utilization
of oxygen by corals is actually due to oxidation of mucus secreted by them during the
course of the experiment. The amount of mucus varies greatly in different genera and may
also be increased at certain times, e.g. during planulation. In view of this source of error it
is impossible to accept their face value figures which claim to represent either the absolute
or the comparative rates of respiration in different corals, or general conclusions which are
based on these figures. Oxidation of mucus may be expected to affect the apparent rate of
respiration in all aquatic animals which normally secrete mucus.
421. Yonge, C. M. and H. M. Nicholas. 1940. Structure and function of the gut and symbiosis with
zooxanthellae in Tridachia crispata (Oerst.) Bgh. Papers Tortugas Laboratory 32: 287-
301 (issued Sept. 1940).
Carnegie Institution of Washington Publication Number 517.
During the visit of the senior author to the Tortugas Laboratory in the season of 1934, a
number of specimens of a very interesting and beautiful species of opisthobranch mollusk
were collected a low tide on the surface of the reefs. Examination revealed the invariable
presence of brown unicellular algae, or zooxanthellae, within their tissues. Tridachia
crispata is an elysoid opisthobranch with the body extremely flattened dorsoventrally and
extended into undulating body folds laterally and terminally. It occurs under stones on the
reefs of the Torugas group and elsewhere in the West Indies. The feeding and digestive
104
systems are described. These have the typical elysoid structure with modifications, notably
in the digestive diverticula, correlated with the excessive flattening of the body. Like the
other members of the Elysiidae, T. crispata is a highly specialized herbivore.
Zooxanthellae are habitually present in a restricted zone a short distance from the margin
of the body fold. They occur freely within the connective tissue and increase by division.
There is no evidence that the animal normally consumes them, but reasons are given for the
suggestion that they may be of value to the animal by removing waste products of
metabolism produced within the body fold .
422. Zeleny, C. 1907. The effect of degree of injury, successive injury, and functional activity upon
regeneration in the scyphomedusan Cassiopea xamachana. Journal of Experimental
Zoology 5: 265-74.
This study is part of a series of experiments at the Dry Tortugas Marine Lab on the internal
factors controlling regeneration in Cassiopea and other forms, including the degree of
injury and successive removal of a part and rhythmical pulsations of the disk. The removal
of 6 to 8 arms constitutes the most favorable degree of injury for the regeneration of each
arm. When comparing the rate of regeneration of disks, where the disk was made to
pulsate rhythmically with cases without pulsation, there is no advantage in favor of the
pulsating ones, but rather a retardation. Other tests of successive injury upon regeneration
were made on chelae of the gulf-weed crab, Portunus sayi, which reveal that the second
regeneration is greater than the first. However, when the age factor is removed the two are
exactly alike.
423.
. 1908. Some internal factors concerned with the regeneration of the chelae of the gulf-weed
crab (Portunus sayi). Papers Tortugas Laboratory 2: 103-38.
Carnegie Institution of Washington Publication Number 103.
The primary object of the experiments described was twofold: the quantitative
determination (1) of the effect of successive removal of an organ upon its power to
regenerate and (2) of the character of the changes, if any, produced in the uninjured parts
of the animal by such removals. It was found that (1) individuals of Portunus sayi with a
cephalo-thoracic length between 3-9 and 14.5 mm. show but a slight correlation between
the length of the molting period and the size or age of the animal. (2) The amount of
regeneration of the right chela between the same limits of size is likewise but slightly
correlated with the length of the molting period, but is very closely correlated with the size
of the animal. (3) The specific amount of regeneration of the right chela increases slightly
with increase in size or age of the animal.(4) The specific length of the left chela in
uninjured individuals increases slightly with increase in size or age of the animal.(5) The
proportion between the amount of regeneration of a chela and the length of the chela in
uninjured individuals of the same size is constant, uninfluenced by the size of the animal.
(6) In single individuals the third regeneration is greater than the second and the second is
greater than the first. (7) When the correction for change in the power of regeneration with
size or age is made, it is found that successive removal neither retards nor accelerates the
regeneration of the right chela. (8) The right chela is slightly larger that the left in a great
majority of the individuals. (9) The removal and regeneration of the right chela produces
no change in the growth of the uninjured left chela.
424. Zheng, W. and E. S. Van Fleet. 1988. Petroleum hydrocarbon contamination in the Dry Tortugas
USA. Marine Pollution Bulletin 19, no. 3: 134-36.
The present study extends a previous work westward to the point where Florida Keys
island chain intersects the Gulf Loop Current. Since the Dry Tortugas are located in this
unusual area, they provide an ideal location for examining the fate of petroleum discharged
into the eastern Gulf of Mexico. Beach tar samples were collected along 1 m wide
transects at 18 stations according to the procedures established by CARIPOL (1980). The
105
distribution of Dry Tortugas beach tar ranged from 0.6 g m super (-2) to 22.1 g m super (-
2) dry weight with an average of 9.2 plus or minus 7.8 g m super (-2). There appear to be
no strong correlations between Dry Tortugas beach tar concentrations and either
predominant wind direction or major Gulf Loop Current circulation patterns.
106
Agassiz, A., 1, 2
Andres, B., A., 3
Applegate, R. D., 298
Aska, D. Y., 327
Austin, O. L. Jr., 4
Awbrey, F. T., 23
Baden, D. G., 333, 375
Bailey, E., 5
Bailey, P. L., 6
Baker, B., 7
Ball, S.C., 8
Ballantine, D. L. 9, 403
Barnes, G. W., 72
Bartsch, P., 10, 11, 12, 13, 14
Battey, J. F., 287
Bellow, T. H., 15, 16, 324
Below, L. C., 315
Bennett, F. M., 17
Berrill, N. J., 18
Blinks, L. R., 19
Bohnsack, J. A., 20
Bortone, S. A., 21
Boschma, H., 22
Bowles, A. E., 23
Bowman, H. H. M., 24
Boyden, A., 25
Bradbury, R. C., 26
Braman, R. S., 154
Bramen, R. S., 356
Breder, C. M. Jr., 27, 28
Brinley, F. J., 29
Brooks, H. K., 30
Brooks, W. K., 31, 32
Brown, D.E.S., 33
Brown, W.Y,. 34
Bullington, W. E., 35
Burkenroad, M., 36
Caira, J. N., 37
Calder, D. R., 38
Carrier, J. C., 39, 40
Cary, L.R., 41- 48
Cate, C.N., 49
Ceurvels, A. R,. 105
Chambers, E. L., 50
Chiang, L., 288
Child, C. A., 51
Clapp, R. B., 52
Author Index
Clark, H. L., 53
Clark, L. B., 54
Cochran, W. W., 298
Cole, L. J., 55
Collie, M. R., 56
Colman, J., 57
Conger, P.S., 58
Conklin, E. G., 59, 60
Coonfield, B. R., 61, 62
Coutiére, H., 63
Cowles, R. P., 64, 65
Criales, M.M., 66
Cubit, J. D., 196
Cushman, J. A., 67, 391
Cutright, P. E., 68
Dall, W. H., 69
Darby, H. H., 70-72
Davis, G. E., 73- 82
Davis, J.H., 325
Davis, J. H. Jr., 83, 84
Davis, R. A. Jr., 85
de Laubenfels, M. W., 86- 88
de Renyi, G. S., 89
Deflaun, M. F., 90
Dinsmore, J. J., 91, 92, 137
Dodrill J. W., 81, 82
Dole, R. B., 93
Domeier, M. L., 94
Donaldson, H. H., 95
Donan, P., 356
Donnelly, K. B., 166-168, 356
Doyle, M.M., 98
Doyle, W. L., 96-98
Drew, G. H., 99
Dustan, P., 100, 101,163
Edmondson, C..H., 102
Erseus, C., 103
Farfante, I. P., 104
Feinstein, A. A., 105
Feltham C. B., 111
Fenimore Johnson, E. R., 72
Ferguson, John C., 301
Field, R. M., 106, 107
Fisk, E. J., 108
Forcucci, D., 264
ECO
Fosberg, F. R., 350
Gardiner, M. S., 369
Gault, C.C., 251
Gee, H., 110, 111
Gersh, I.. 112
Gilmore, R. G., 113
Ginsburg, R.N., 114
Given, B., 316
Goldfarb, A. J., 115-122
Goldman, M. A., 391
Goodrich, H. B., 123
Gordon, M., 124
Goy, J. W., 125
Grave, C., 126-128
Gudger, E. W., 129-133
Haight, F. J., 374
Halas, J.C., 101
Halley, R.B., 134, 328, 330
Hanlon, R. T., 135
Hargitt, C. W., 136
Harper, D. E., 20
Harrington, B. A., 137
Harris, J.E., 28, 138
Harrison, C.S., 354
Hartman, C.G., 139
Hartmeyer, R., 140, 141
Harvey, E.N., 142-145
Hatai, S., 146-148
Hayes, F.R., 149, 150
Heard, R. W., 151
Helwig, E.R., 152
Hendee, E. C., 153
Hendrix, S.A. 154
Hess, W.N., 54, 155, 156
Hildebrand, S. F., 211, 212
Hine, A. C., 203
Hixon, R. F., 135
Hoetjes, P.C., 400
Hoffman, W., 157, 411
Holmes, C. W., 158
Hooker, D., 159
Hopkins, D. L., 160
Howe, M.A., 391
Hudson, J. H., 310, 328, 330
Hutton, R.F., 105
Jaap, W.C., 101, 161-168, 264, 356, 403
Jacobs, M.H., 169
107
Jefferson, J. P., 170
Jehl, J. R., 23
Jennings, H.S., 171
Jindrich, V., 172
Jones, N., 173, 285
Jones, R. D., 174
Jones, R.S., 113,175
Jordan, D. S., 176, 177
Jordan, H. E., 178-186
Kaas, P., 187
Kale, H. W., 188
Keiller, V.H., 369
Kellner C., 32, 189
Kille, F. R., 190
Kilma, E. F., 261
Kindinger, J. L., 330
King, J. Jr., 276, 277
Kojis, B. J., 167, 168, 403
Kopac, M. J., 191
Kunkel, B. W., 192
Lashley, K.S., 398
Le Compte, M., 193
Lee, T.N., 66
Leitch, J.L., 194, 195
Lessios, H. A., 196
Lidz, B. H., 328, 330
Linton, E., 197-199
Lipman, C. B., 200- 202
Locker, S. D., 203
Longley, W. H., 204-212
Lucké, B., 213-215
Lynts, G. W., 216
Lyons, W.G., 163, 217
Mann, A., 218, 374
Manter, H. W., 219-221
Marsh, G., 222- 224
Martin, D. F., 356
Martin, J.C., 346
Martin, L. K., 40
Mason, C.R., 316, 318
Mast, S. O., 225
Matthai, G., 226
Maul, George A., 394
Mayer, A. G., 227-242
McClellan, D.B., 20
McClendon, J. F., 243-251
McKenna, J. E. Jr., 167, 168, 403
108
Meeden Jars) 252
Meier, O. W., 288
Meyers, P.A., 401
Miller, H. M. Jr., 253
Miller, R. A., 254
Milligan, M.R., 103
Millspaugh, C. F., 255
Mitchell-Tapping, H. J., 256
Moore, T., 170
Moritz, C. E., 257
Mulholland, S., 251
Multer, H. G., 258, 259
Mumme, R. L., 415
Murphy, L. E., 260
Nance, James M., 261
Nicholas, H.M., 421
-Nicoll, P. A., 128
O'Neill, C. W., 85, 262, 263
Ogden, J.C., 264
Osburn, R. C. , 265
Patella, F.J., 261
EV Iiy, 12 (C IS, S55)
Paul, J.R., 281
Paulson, D.R., 318
Payne, F., 266, 267
Pearse, A. S., 268-274
Perkins, H. F., 275
Perlmutter, D.G., 151
Petrovic, C. A., 276, 277
Phillips, A. H., 278-280
Pichard, S. L., 281
Pitts, R. F., 282
Plan Development Team, Reef Fish
Management
Plan, South Atlantic Fishery Management
Council., 283
Plantier, T. L., 284
Plough, H. H., 285
Porter, J. W., 264, 286-288
Porter, J. Y., 170
Potthoff, T., 289
Powers, P. B. A., 290, 291
Pratt, H. L. Jr., 39, 40
Pratt, H. S., 292- 297
Pritchard, M.H., 37
Raim, A., 298
Rebenack P., 21
Reighard, J., 299, 300
Reynolds, J. E. II, 301, 302
Richards, O. W., 303, 336
Richards, W. J., 289
Richardson, T., 288
Ricklefs, R. E., 304, 305, 405
Riley, G. A., 306
Riska, D. E., 307, 308
Rivas, L. R., 309
Roberts, H H., 310, 311
Robertson, D.R., 196
Robertson, W. B. Jr., 4, 5, 34, 52, 92, 157, 312-
318, 324, 355, 405, 411, 413-416
Robinson, A. H., 319
Rouse, L. J. Jr., 310, 311
Sarver, S. K., 332
Schaeffer, A. A., 320, 321
Schlumberger, H.G., 215
Schmahl, G. P., 403
Schmidt, T. W., 373
Schmitt, W. L., 322
Schnell, G. D., 323
Schreiber, R. W., 324
Schroeder, P. B,. 325, 365
Scott, W. E. D., 326
Seaman, W., Jr., 327
Shimizu, Y., 333
Shinn, E. A., 203, 328-330
Shoemaker, C. R., 331
Siegel, D. M.., 21
Silberman, J. D., 332
Smayda, T. J., 333
Smith, G. J., 287
Smith, H. B., 254
Smith, H. G., 334
Smith, N. P., 264
Snoek E., 105
South Florida Area Study 335
Spence, J., 336
Sprunt, A. Jr., 337-343
Steinen, R. P., 134
Steinmetz, J. C., 302
Stevenson, J. O., 344
Steward, F. C., 345, 346
Stockard, C. R., 347-349
Stoddart, D. R., 350
Stone, R. G., 351-353
109
Stoneburner, D. L., 354, 355 Yamanouchi, S., 417
Strom, R. N., 356 Yonge, C. M., 418-421
Stromsten, F. A., 357, 358
Szmant, A. M., 264
Zeleny, C., 422, 423
Zheng, W., 424
Tandy, G., 359
Tartar, V., 360
Tashiro, S., 361
Taylor, C. V., 371
Taylor, Jane B., 362
Taylor, W. R., 363, 364
Teas, H. J., 365
Tennent, D. H., 366-371
Thiele, J., 372
Thompson, J.C., 177
Thompson, M. J., 175, 373
Thorp, E. M., 374
Tomas, C. R., 333, 375
Torrey, H. B., 376
Treadwell, A. L., 377-379
Ubelacker, J. M., 380
Van Fleet, E.S., 424
Vaughan, T. W., 374, 381-391
Visscher, J. P., 392
Vukovich, F. M., 393, 394
Walker, N. D., 310, 311
Wallace, W.S., 395
Walsh, P.J., 332
Wartman, W., 396
Watson, J. B., 397, 398
Wells, R.rC., 399
Westinga, E., 400
Westrum, B.L., 401
Wheaton, J., 164-168, 402, 403
Whitaker, D. M., 371, 404
White, S.C. 305, 405, 415
White-Schuler, S. C., 304
Wichterman, R., 406, 407
Williams, O. L., 408
Willier, B. H., 409
Wilson, C. B., 410
Winegarner, C. E., 15, 411
Wolfe, C. A., 412
Woolfenden, G.E., 4,5, 413-416
110
Subject Index
Accipiter striatus, 3
Acropora, 241
Acropora cervicornus, 30, 73, 74, 193
Acropora palmata, 74, 172, 328, 388
Actinian, 60, 136
Alcyomeum, 48
Alcyonaria, 48, 407
Alpheidae, 63, 70, 248
Alpheus, 63, 70
Amphipods, 151, 271, 331
Anchor damage, 73
Animal succession, 57
Anisonema vitrea, 102
Annelids, 119; 120, 228, 351, 352, 353, 377,
378, 379, 380
Anous stolidus, 34, 307, 308, 405, 412
Ants, 55, 242
Aplopus, 178, 186, 347
Aplysia protea, 257
Ascidians, 18, 126, 127, 128, 140, 228, 285,
303,
336, 376
Asio flammeus, 157
Astroea longispina, 43
Astropecten duplicatus, 65
Atlantic palolo (Worm), 227, 228
Atmospheric compounds, 154, 249
Audouinella ophioglossa, 9
Audubon, 10
Avicennia, 83
Banded red knot, 413
Barnacles, 392
Bathydrilus oligochaeta, 103
Beach rock, 106, 107, 114, 259
Beach tar, 424
Belonidae, 27
Bioluminescence, 144, 145
Bird history, 338, 339, 343, 344, 397
Birds, 4, 5, 10, 17, 108, 137, 157, 316, 326, 335,
337, 338, 339, 341, 342, 344, 413
Black noddy, 313, 318
Black phoebe, 108
Black water event, 163, 170
Blood relationships, 25
Bluefin tuna, 289
Botanical ecology, 24
Brackish-water ponds, 268
Breeding, 11, 13, 14, 284, 308
Briareum, 48, 163
Brown anole, 411
Brown noddy, 34, 305, 307, 308, 313, 397, 398,
405, 412
Brown pelican, 15, 324
Bryozoans, 265
Bubulcus ibis, 415
Calcium carbonate, 99, 106, 107, 201, 202, 246,
249, 259, 274, 385, 391
Calidris canutus, 413
Cardinal, 16
Cardinal cardinalis, 16
Cassiopea, 44, 45, 47, 142, 228, 237, 244, 245,
334
Cassiopea xamachana, 116, 146, 147, 148, 231,
234, 235, 238, 275, 334, 349, 422
Cattle egrets, 137, 415
Caulerpa, 417
Cerions, 11, 12, 13, 14
Chart, 109
Ciliates, 35, 290, 291, 360, 406, 407
Clibanarius, 192
Cnideria, 22, 41, 42, 46, 48, 59, 98, 135, 146,
147, 148, 161, 162, 168, 171, 172, 193, 228,
231, 234, 235, 237, 241, 244, 245, 334, 349,
395, 402, 403, 418, 419, 422
Coelenterates (Cnideria), 144, 275
Color patterns, 61, 62, 123, 130, 204, 205, 206,
207, 209, 210, 299
Common tern, 304
Condylactis gigantea, 135
Conocarpus, 83
Coral bleaching, 161, 288
Coral growth, 335, 385, 388
Coral mortality, 287, 288, 310, 311
Coral polyps, 22
Coral reef, 1, 2,57, 73, 161, 162,163; 16s-steve
168, 252, 264, 287, 288, 310, 311, 359, 381,
382, 383, 384, 385, 386, 387, 390, 403
Coral reef sampling, 165, 167, 288, 359, 403
Coral reef structure, 74, 79, 100, 168, 172, 335,
385, 387, 389, 391
Corals, 98, 226, 229, 241, 248, 252, 287, 383,
402, 403, 418, 420
Crabs, 33, 71, 192, 204, 257, 274, 303, 422, 423
Crangon armillatus, 62, 70, 71, 89, 156, 257
Crustaceans, 66, 71, 75, 78, 81, 89, 104, 125,
145, 151, 156, 192, 204, 248, 261, 274, 303,
322, 331, 410, 422, 423
Cultural resources, 260
Currents, 384, 389, 393, 394, 424
Cytology, 33, 43, 50, 59, 87, 96, 97, 98, 117,
SAT I2 NAS 49 ISON IS2 153578;
179, 180, 181, 182, 183, 184, 185, 186, 191,
194, 195, 222, 223, 224, 244, 247, 254, 257,
266, 267, 320, 321, 336, 345, 346, 349, 353,
360, 366, 367, 368, 370, 371, 376, 404, 406,
409, 417
Diplectrum vittatum, 21
Diadema, 196
Diatoms, 58, 218
Diplectrum formosum, 21
Diporia clivosa, 168
Diving helmet, 131
DNA,90, 332
Dove, 315
Echinaster, 65, 182
Echinoderms, 53, 118, 121, 122, 149, 150, 153,
182, 185, 190, 194, 195, 196, 247, 254, 280,
290, 291, 360, 366, 368, 369, 370, 371, 376,
404
Echinometra lacunter, 53, 150, 194, 195, 254,
Sil, SHG)
Ecology, 83, 84
Effect of light on organisms, 28, 65, 72, 155,
156, 159, 222, 223, 334, 346, 368
Effect of photosynthesis, 230, 246
Effects of radiation, 351, 352
Effects of temperature on organisms, 224, 229,
234, 241, 244, 287, 310, 311, 388, 390, 393,
394
Empidonomus varius, 26
Epinephelus, 282
Epinephelus morio, 273
Evolution, 138
Exococetidae, 27
Fish, 21, 27, 28, 29, 33, 36, 39; 61, 68, 94, 112,
MB IS 24 29M IG OMS2lS3 135385
139, 170, 176, 177, 205, 206, 207, 209, 210,
DEO NSS ANA S22 1295 282,283;
289, 299, 309, 373, 408, 410
Fish communities, 175, 212, 283, 373
Fish kill, 310
Fish sampling techniques, 175, 205
Fish tumors, 213, 214, 215
Flight speed (Birds), 323
Florida Keys National Marine Sanctuary, 7, 20
Food habits, 21, 92, 171, 205, 397, 421
Foraminiferae, 67, 216
Frigate birds, 323
Gastropods, 89
LU
Geologic history, 381, 382, 383
Geology, 30, 83, 106, 107, 114, 134, 172, 203,
256, 258, 259, 260, 262, 263, 302, 328, 329,
374, 381, 382, 383, 385, 391
Ginglymostoma cirratum, 39, 112, 129, 133
Gorgonacae, 41, 42, 46, 402
Gorgonia, 48, 402
Great black-backed gull, 188
Growth, 303, 304, 305, 346, 348, 405,
412
Habits, 64, 132, 133, 248, 272, 273,
275, 347, 398, 418, 419, 421
Haemulidae, 36
Halichores, 213
Halichores bivittatus, 124
Halimeda, 263
Halimeda hummii, 9
Hatching success, 23
Hemirhamphidae, 27
Histology, 190
Holothuria, 190, 279, 280, 369
Homing instinct, 10, 398
House sparrow, 416
Hydroids, 38, 116, 376, 395
Hydrology, 134
Hypoglossum rhizophorum, 9
Hypoplectrus, 94, 373
Insects, 8, 347
Invertebrates, 6, 8, 11, 12, 13, 14, 18, 22, 25, 32,
35, 37, 38, 41, 42, 43, 44, 45, 46, 47, 48, 49,
50, 51, 53, 54, 55, 58, 59, 60, 62, 63, 64, 65,
66, 67, 69, 70, 71, 73, 74, 75, 76, 77, 78, 80,
81, 82, 86, 87, 88, 89, 98, 100, 101, 102, 103,
104, 117, 118, 119, 120, 121, 122, 125, 126,
127, 128, 135, 136, 140, 142, 143, 144, 145,
146, 147, 148, 149, 151, 152,.153, 155, 156,
161, 162, 169, 171, 172, 178, 182, 183, 185,
186, 187, 189, 190, 192, 193, 195, 196, 197,
Ie, ID, Bil, PIO, QAO, PA, P27, Q29), Z3i-
235, 238, 239, 242, 244, 245, 246, 247, 248,
253, 254, 261, 265, 266, 267, 270, 271, 272,
275, 279, 280, 285, 290, 291, 295, 294, 295,
296, 297, 303, 322, 331, 334, 347, 351, 352,
353, 360, 366, 367, 368, 369, 376, 377, 378,
379, 380, 392, 395, 400, 402, 408, 418, 419,
421, 422
Iotrochota, 87, 152
Ischnochiton, 187, 217
Land-birds, 3, 16, 26, 276, 277, 315, 317, 326,
341, 415, 416
112
Larval shrimp, 66
Launcularia, 83
Least terns, 414
Leodice fucata, 54
Limulus, 184, 228
Linckia, 360
Lipogramma anabantoides, 113
Littoral, 269, 272
Loggerhead turtle, 159, 179, 180, 181, 183, 198,
295, 297, 357, 358, 409
Long Term Ecological Research (LTER) 163,
167, 225, 264
Longevity, 34
Lutjanus griseus, 124, 214
Lytechinus variegatus, 50, 153, 247, 368, 370,
404
Maeandra aerolata, 22, 229, 241, 418
Maps, 1, 74
Manatees, 301
Mangrove, 83, 325, 365
Marine algae, 9, 19, 96, 97, 191, 222, 223, 224,
345, 346, 363, 364, 417
Marine amoebe, 160, 320, 321
Marine bacteria, 99, 110, 111, 202
Marine fishery reserves (MFR), 283
Marine Laboratory, 232, 236, 240
Marine parks, 79, 319
Masked booby, 52
Mastogloia, 218
Mating, 39
Medusae, 44, 45, 228, 231, 234, 239, 275, 349
Merlin, 298
Metals in organisms, 278, 279, 280, 354, 355,
356
Migration, 17
Millepora complanata, 161, 172
Mites, 274
Mollusks, 11, 12, 13, 14, 49, 69, 89, 187, 217,
DIRE) eed, 92, C19) |
Monocotyle, 292
Monospotus indicus, 9
Montastrea annularis, 162, 168, 172, 241, 385
Montastrea cavernosa, 168
Morphodynamics, 85, 192
Mosquitoes, 8
Myliobatis, 292
Nematodes, 408
Neoplastic growths, 124
Nesting, 15, 324, 343, 397, 414, 416
Nurse shark, 39, 40 112, 129, 133, 197
Ocypoda, 64
Oikopleura, 189
Oil spill, 335
Ophicoma, 182
Orbicella (Madrepora), 193, 229, 241, 388
Osmotic pressure, 269
Owl, 157
Panulirus argus, 75, 77, 78, 89, 156, 327, 332
Parasitic copepods, 410
Parasitic isopods, 273, 410
Parasitic worms, 37, 197, 198, 199, 219, 220,
221, 253, 292, 293, 294, 295, 296, 297
Pedibothrium, 37
Penaeus duorarum, 66, 261
Pesticides, 356
PH, 230, 250, 251, 334
Physiology, 47, 97, 111, 135, 160, 169, 334,
345, 346, 357, 358
Phytoplankton, 333, 375
Plankton, 306
Plectognath, 28
Plexaura, 46
Pollution, 335, 424
Polycistor, 140
Polyplacophora, 217
Pomacentrus, 29
Pomacentrus leucostictus, 61
Porites, 229, 241
Porites asteroides, 168
Portunus sayi, 422, 423
Promicrops (Epinephelus) itajara, 273
Protozoan, 102, 160, 169, 406, 407
Pseudocyphoma, 49
Ptychodera, 43, 89, 155, 266, 267
Pycnogonida, 51
Rails, 277
Rat, 95
Recreational headboat fishery, 20
Red-headed woodpecker, 317
Red tide, 105
Reef fishes, 20
Reef formation, 2, 41, 42
Reefs, 30
Regeneration, 6, 44, 45, 47, 115, 116, 120, 152,
190, 235, 285, 348, 349, 351, 352, 353, 360,
422, 423
Replenishment reserves, 7, 20, 283
Rhizophora, 83
Richardina spinicincta, 125
Roseate terns, 313, 414
Sabellids, 6, 352, 379
Sabines's gull, 56
Salinity, 93, 116
Salpa floridian, 31
Scleractinia, 161
Scyphomedusa (jellyfish), 59, 234
Sea-birds, 4, 15, 23, 34, 52, 56, 91, 92, 188,
284,
298, 304, 305, 307, 308, 312, 313, 318, 323,
324, 340, 341, 354, 355, 397, 398, 405, 412,
414
Sea-cucumber, 190
Sea-level change, 203, 258, 262, 263
Sea turtles, 159
Sea-urchin, 50, 117, 121, 122, 150, 153,
194, 195, 196, 247, 254, 290, 291, 366,
368, 370, 371, 404
Sea-water composition, 72, 90, 93, 115,
174, 200, 201, 202, 230, 235, 237, 238,
243, 246, 249, 250, 251, 281, 320, 361,
389, 399, 401
SEAKEYS, 264
Sedimentation, 85, 172, 256, 258, 259,
262, 263, 329, 374
Sharks, 39, 139
Sharp-shinned hawks, 3
Sicyonia penaeoidea, 104
Siderastera, 168, 229, 241, 385, 419
Snapping shrimps, 63, 322
Sooty terns, 4, 5, 23, 91, 92, 284, 304,
312, 313, 314, 354, 355, 397, 398, 405
Speculata advena, 49
Spheciospongia vesparia, 151, 400
Sphyraena barracuda, 132
Spiny lobster, 75, 76, 77, 78, 80, 81, 82,
ISO, S27, Sw
Sponges, 86, 87, 88, 101, 151, 152, 270, 400,
403
Sport harvest, 75, 76, 78, 79, 81, 82
Starfish, 65, 360, 369
Starvation, 231, 334, 415
Stephanochasmus, 294
Sterna fuscata, 4, 5, 91, 92, 284, 304, 314, 354,
405
Sterna hirundo, 304
Sterna melanoptera, 340
Sting ray, 68
Storms, 17, 405
Sula dactylata, 52
Syllids, 380
Synentognathi, 27
113
Topography, 1, 84, 325, 328, 329, 343, 350
Transatlantic migration, 314
Transplanting organisms, 403
Trematodes, 220, 221, 253, 292, 293, 294, 295,
296, 297
Trichechus, 301
Tunicata, 31, 32
Valonia, 19, 96, 97, 191, 222, 223, 224, 345,
346
Vanadium, 279, 280
Variegated flycatcher, 26
Vegetation, 24, 84, 255, 325, 350, 365
Vessel groundings, 335, 403
Vocal signals, 307, 308
Worms, 6, 54, 103, 119, 120, 155, 197, 198,
DY, By ALVZ, Psa, 22), P23)5 P2O, PM, SIS,
353, 377, 378, 379, 380, 408
Zooxanthellae, 98, 334, 407, 421
*# U.S. GOVERNMENT PRINTING OFFICE: 1998- 434-814
ATOLL RESEARCH BULLETIN
NOS. 443-449
NO.
NO.
NO.
NO.
NO.
NO.
NO.
443.
444.
445.
446.
447.
448.
449.
THE EVOLUTION OF A HOLOCENE FRINGING REEF AND
ISLAND: REEFAL ENVIRONMENTAL SEQUENCE AND SEA
LEVEL CHANGE IN TONAKI ISLAND, THE CENTRAL
RYUKYUS
BY H. KAN, N. HORI, T. KAWANA, T. KAIGARA, AND K.
ICHIKAWA
CHECKLIST OF THE SHOREFISHES OF OUVEA ATOLL, NEW
CALEDONIA
BY MICHEL KULBICKI AND JEFFREY T. WILLIAMS
ON THE ORIGIN OF DRIFT MATERIALS IN THE MARSHALL
ISLANDS
BY D.H.R. SPENNEMANN
DISTRIBUTION OF RAT SPECIES (RATTUS SPP.) ON THE
ATOLLS OF THE MARSHALL ISLANDS: PAST AND PRESENT
DISPERSAL
BY D.H.R. SPENNEMANN
A POSSIBLE LINK BETWEEN CORAL DISEASES AND A
CORALLIVOROUS SNAIL (DRUPELLA CORNUS) OUTBREAK IN
THE RED SEA
BY ARNFRIED ANTONIUS AND BERNHARD RIEGL
MARINE ALGAE FROM OCEANIC ATOLLS IN THE
SOUTHWESTERN CARIBBEAN (ALBUQUERQUE CAYS,
COURTOWN CAYS, SERRANA BANK, AND RONCADOR BANK)
_BY GUILLERMO DIAZ-PULIDO AND GERMAN BULA-MEYER
SCIENTIFIC STUDIES ON DRY TORTUGAS NATIONAL PARK:
AN ANNOTATED BIBLIOGRAPHY
BY T.W. SCHMIDT AND L. PIKULA
ISSUED BY
NATIONAL MUSEUM OF NATURAL. HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
OCTOBER 1997
HECKMAN ll
BINDERY INC. [4
JUNE 98
‘To-Plea® N. MANCHESTER,
Bound -To-Pleas? yA 46962
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